Earliest Porotic Hyperostosis on a 1.5-Million-Year-Old Hominin,Olduvai Gorge,Tanzania

Meat-eating was an important factor affecting early hominin brain expansion, social organization and geographic movement. Stone tool butchery marks on ungulate fossils in several African archaeological assemblages demonstrate a significant level of carnivory by Pleistocene hominins, but the discovery at Olduvai Gorge of a child''s pathological cranial fragments indicates that some hominins probably experienced scarcity of animal foods during various stages of their life histories. The child''s parietal fragments, excavated from 1.5-million-year-old sediments, show porotic hyperostosis, a pathology associated with anemia. Nutritional deficiencies, including anemia, are most common at weaning, when children lose passive immunity received through their mothers'' milk. Our results suggest, alternatively, that (1) the developmentally disruptive potential of weaning reached far beyond sedentary Holocene food-producing societies and into the early Pleistocene, or that (2) a hominin mother''s meat-deficient diet negatively altered the nutritional content of her breast milk to the extent that her nursing child ultimately died from malnourishment. Either way, this discovery highlights that by at least 1.5 million years ago early human physiology was already adapted to a diet that included the regular consumption of meat.     

Late Pliocene grassland from Olduvai Gorge,Tanzania

The Olduvai fossil plants documented by us in this paper are the first direct evidence for open grassland in the late Neogene of Africa based on macroplant remains. Silicified remains of herbaceous ground cover are exceptionally well preserved in situ within Late Pliocene sediments below the initial pyroclastic ash surge unit of Tuff IF in the uppermost part of Bed I, Olduvai Gorge, northern Tanzania. Published radiometric and palaeomagnetic dates place this grass layer between 1.839 + 0.005 Ma and 1.785 + 0.01 Ma. Exposed at localities on the south side of the Gorge this herbaceous ground cover grew on a floodplain developed on a dried out lake bed, following pronounced lake retreat of saline–alkaline palaeo-Lake Olduvai during a developing dry climatic phase. Sheathed basal culms, rhizomes and roots are interpreted as those of one or more small mat-forming grasses or less likely, sedges. Small dicotyledonous herbs were probably also present. The proximity of adjacent plants indicates a relatively dense ground cover. Roots extended at least 8 cm below the ground surface. Aerial parts of the plants were absent or were not preserved when the weathered basal culms were covered by a thin layer of brown waxy clay, followed by fallout of pyroclastic ash. Both units were mostly eroded away prior to emplacement of a wet, cool pyroclastic surge which then buried and preserved in situ remnants of the herbaceous ground cover. Preservation of the semi-woody rhizomes implies well-drained soils, otherwise the plant material would have quickly rotted. Collections from discontinuous exposures indicate the grassland covered an area of at least a few hectares. This open grassland would have provided grazing for the Late Pliocene fauna.     

Middle Pleistocene hominids from Olduvai Gorge, northern Tanzania

Cranial, dental, and mandibular remains of eight Olduvai hominids are described in detail. Four individuals were recovered in situ in Beds II to IV, while three more are most probably derived from Bed IV, the Masek Beds and the Lower Ndutu Beds. One specimen is of uncertain provenance. Deposits from which the fossils were collected range from late Lower Pleistocene to Middle Pleistocene in age. Of particular interest are three fragmentary lower jaws, which can be compared to mandibles of Homo erectus known from localities in Northwest Africa and China. Olduvai hominid 22, a nearly complete half mandible with crowns of P3-M2 in place, shares many anatomical features with fossils from Ternifine and Choukoutien. This individual is also similar to a jaw from the Kapthurin Formation west of Lake Baringo, Kenya. How best to interpret these comparisons is not clear, but in view of marked similarities between specimens representing geographically diverse populations from different time periods, it may be unwise to rely on mandibular evidence alone to document the presence of regional lineages. Gradual change and continuity within a sequence of Northwest African Homo fossils has been endorsed by many workers, but such hypotheses cannot be tested adequately with the fragmentary jaws available.     

Taphonomic perspectives on hominid site use and foraging strategies during Bed II times at Olduvai Gorge, Tanzania

The faunal assemblages excavated by Mary Leakey in Bed II of Olduvai Gorge, Tanzania, have, like the more well-known Bed I assemblages, traditionally been interpreted as the result of hominid butchering activities in the lake margin and riverine settings of the paleo-Olduvai Basin. A reexamination of all of Leakey's Bed I sites has shown that hominids played little or no role in the formation of all but one of those faunal assemblages, a finding that prompted the reanalysis of the Bed II sites presented here. We expand upon a previous taphonomic study that provided systematic data for HWK East Levels 1–2, MNK Main, and BK. In addition to these assemblages, we provide data on HWK East Levels 3–5, FC West, TK, and SHK. Our data contradict previous interpretations of MNK Main as a hominid accumulation but uphold the contention that BK represents a primarily hominid accumulation reflecting early access to carcasses. The small and poorly preserved assemblages from FC West and TK are difficult to link unambiguously to either hominids or carnivores. Site MNK Main and HWK East Levels 3–5 appear to be death arenas where carcasses accumulated via natural deaths and/or serial predation. Site SHK is severely biased by selective retention and therefore little can be said of its formational history. Nevertheless, no hominid modifications were documented in this assemblage. Comparisons with other Olduvai sites indicate a more conspicuous hyena taphonomic signal during Bed II times than Bed I times, which appears to mirror the changing configuration of the large carnivore guild. These findings also beg the question of what activities were being carried out by hominids with the stone tools discarded at these sites. Although it seems clear that hominids were utilizing stone tools to carry out subsistence activities unrelated to carcass butchery, more excavation and techniques such as phytolith analysis should be employed to explore alternative explanations.     

Tracks,Trails and Trampling by Large Vertebrates in a Rift Valley Paleo-Wetland,Lowermost Bed II,Olduvai Gorge,Tanzania

The impact of large vertebrates on the sedimentary record in an East African groundwater-fed wetland in Ngorongoro Crater, Tanzania, is used as a modern analog to interpret an ancient (Plio-Pleistocene) wetland record in Olduvai Gorge, Tanzania. The 3 km² wetland at Olduvai is characterized by massive silty claystones produced by intense vertebrate trampling, as well as trails by vertebrates frequenting the wetlands and isolated footprints that represent a “snapshot in time.” Based on modern analogs, a paleo hippo trail (1.2 m wide and 0.6 m deep) in the Plio-Pleistocene wetlands is interpreted as a frequently used corridor between hippo pools (days) and grazing meadows (nights). Groundwater-fed wetlands are low energy environments where the physical record appears to be dominated by plant and animal activity. The bioturbation record reflects a number of interacting factors such as substrate texture, moisture content, sedimentation rate, frequency of flooding, type of animals present, trampling rate, and post-depositional changes (compaction). Lithofacies in both the modern and ancient wetlands include muddy sandstone (drainage channels) and silty claystone (vegetated and nonvegetated mud flats). Organic-rich sediments eventually oxidize, eliminating most evidence of the habitat. Modern wetlands have organic-rich mud and peat, whereas the ancient analog has siliceous earthy claystones that contains plant remains, bone fragments, pollen, phytoliths, and localized beds of diatomite. Thus, the physical record of vertebrate bioturbation in conjunction with paleontological and lithological records provides crucial information on the ecology of ancient wetland environments.     

Tool-using strategies by early hominids at bed II, Olduvai Gorge, Tanzania

This study examines the current prevailing model of Oldowan technology-the opportunistic, least-effort strategy of stone tool making and using by early hominids. The sample includes the MNK chert factory site and three contemporaneous assemblages from Olduvai Gorge, all dated between 1.65 and 1.53 m.y.a. The analysis suggests that early hominids at Olduvai may have been selective, applying distinctive strategies in making and using tools depending on the different types of raw materials available to them. The preponderance of lava cores and near absence of flakes associated with the cores suggest that lava cores at Olduvai did not provide a source of flakes. They were primarily heavy-duty core tools, despite the fact that the majority of Olduvai lava is of excellent quality for flaking. Contrary to this pattern, the abundance of chert flakes and the lack of large chert cores suggest that the production of flakes was the most important strategy applied to chert. Original forms and flaking mechanics of the raw materials may have been important factors in the simultaneous application of the different, complementary strategies. The Oldowan tool-using strategy was dynamic and flexible, in response to changes in raw material availability. The use of chert between 1.65 and 1.53 m.y.a. was apparently related to the drastic decrease in flake production in lava and quartz. Finally, lack of initial reduction episodes of lava material challenges the idea of the stone cache strategy at Olduvai between 1.65 to 1.53 m.y.a.     

Fossil sedges, macroplants, and roots from Olduvai Gorge, Tanzania

A variety of macroplants has been recorded and collected from the eastern paleolake margin of Olduvai Gorge, Tanzania, from Upper Bed I and Lower Bed II, dated at ∼1.7-1.85 Ma. The plant groups represented are sedges, grasses, and woody and herbaceous dicotyledons. Most of these plants are fragmented, but the roots are in situ. The modes and quality of preservation, however, are very variable. Silicification is the dominant type of preservation; it ranges from high quality faithful replacement of cells resulting in silicified wood and sedge culms that are identifiable on the basis of their internal anatomy, to poor quality biotubes lacking internal anatomy or external features that prevent assignment to a specific plant or invertebrate origin. In between this range are silicified roots and grass culms identified by their external anatomy, and leaf and stem impressions. Interpretation of the paleoecology is limited by the quality of preservation. The in situ root horizons are useful for recognizing paleo-surfaces. The best quality preservation where internal anatomy is preserved occurs at HWK E and MCK, localities that are in the middle of the fault compartments so the vegetation can be reconstructed for these sites. Some sedge culms are described, illustrated, and identified as possible species of Cyperus, Fuirena, and Schoenoplectus.     

Soricidae (Mammalia) from the Early Pleistocene of Olduvai Gorge, Tanzania

Soricid remains collected from Bed I of Olduvai Gorge are described. The great majority of the specimens are mandibles. A survey of the mandibles of living African species revealed many differences in characters of the lower teeth and jaw that can be used for identification. On the basis of these characters, nine species are distinguished in the Olduvai collection, of which six are well enough preserved to permit a discussion of their relationships to living species. Three new species and one new subspecies are described. All the Olduvai shrews differ in some respects from their nearest living relatives; three species are close to species from Makapansgat, Swartkrans and Sterkfontein, RSA, though there appear to be slight differences. A change in the soricid fauna takes place within Bed I, interpreted as due to increasing aridity.     

Fine resolution of early hominin time, Beds I and II, Olduvai Gorge, Tanzania

Reconstructing paleoenvironments and landscapes within lake-centered, hominin-yielding basinal sequences requires a resolution of time-rock units finer than but complementary to that provided by the present tephrostratigraphy. Although indispensable in providing an absolute time frame at Olduvai, the average 15,000-20,000 year intervals between successive tuff units lack the time resolution to construct a sufficiently contemporary paleolandscape within sedimentary intervals away from the interleaved tuffs. Such control is essential to construct valid paleogeographies in which to contextualize contemporaneous paleoanthropological sites and the traces of hominin land use they contain. Within Beds I and II of the Olduvai Basin a Sequence Stratigraphic analysis has achieved a relative time framework in which time-rock units, “lake-parasequences,” each generated by a major advance and withdrawal of the lake system, are recognizable for average periods of about 4000 years duration. Within each of these time slices at least two paleogeographic landscapes are identifiable, reducing the time constraints of an individual landscape reconstruction to a few thousand years. Within the sedimentary succession both highly incised and less incised unconformities are identifiable to provide sequence boundaries. Within each sequence the higher frequency lake-parasequences can be identified by (1) a disconformable base, (2) accretion of sediment during lake transgression and at maximum, (3) a disconformable top caused by lake withdrawal, and (4) a soil profile generated beneath that disconformable land surface. Individual lake-parasequences can be recognized in lake marginal and fan settings, and their imprint can also be seen in the lake setting where, for example, maximum flooding might be marked by a layer of dolomite. Lower Bed II parasequences represent time intervals of <5000 years, while parasequential periods between Tuffs IB and IC in Bed I are <4300 years. Analogous Holocene lake-level changes of the same order in East Africa have a period close to 4200 years. The estimated period is close to that defined by Stadial/Interstadial Dansgaard-Oeschger Events recorded in the Greenland Ice record, which force cycles of period similar to lake-parasequences, both in the Arabian Sea and Lake Malawi. Lake-parasequences not only aid construction of landscapes, they also allow contextualization of individual paleoanthropological occurrences. For instance, a parasequence lies between Leakey’s Level 1 and her butchered Deinotherium occurrence at FLK N. However, elephantid and giraffid skeletons associated with stone artifacts at VEK, uncovered by OLAPP excavations, are situated on the same land surface as a possibly butchered rhinocerid at KK. To complement the existing absolute radiometric time framework, relative Sequence Stratigraphic techniques might be applied to any lake-centered, hominin-yielding basinal sequence, not only those found within East Africa. Because they are climatically controlled, and might plausibly be related to globally driven Dansgaard-Oeschger Events, lake-parasequences and their associated sequences might be correlatable between various East African basins in the Plio-Pleistocene in the same way as they presently are for the Holocene.     

Quantifying Oldowan Stone Tool Production at Olduvai Gorge,Tanzania

Recent research suggests that variation exists among and between Oldowan stone tool assemblages. Oldowan variation might represent differential constraints on raw materials used to produce these stone implements. Alternatively, variation among Oldowan assemblages could represent different methods that Oldowan producing hominins utilized to produce these lithic implements. Identifying differential patterns of stone tool production within the Oldowan has implications for assessing how stone tool technology evolved, how traditions of lithic production might have been culturally transmitted, and for defining the timing and scope of these evolutionary events. At present there is no null model to predict what morphological variation in the Oldowan should look like. Without such a model, quantifying whether Oldowan assemblages vary due to raw material constraints or whether they vary due to differences in production technique is not possible. This research establishes a null model for Oldowan lithic artifact morphological variation. To establish these expectations this research 1) models the expected range of variation through large scale reduction experiments, 2) develops an algorithm to categorize archaeological flakes based on how they are produced, and 3) statistically assesses the methods of production behavior used by Oldowan producing hominins at the site of DK from Olduvai Gorge, Tanzania via the experimental model. Results indicate that a subset of quartzite flakes deviate from the null expectations in a manner that demonstrates efficiency in flake manufacture, while some basalt flakes deviate from null expectations in a manner that demonstrates inefficiency in flake manufacture. The simultaneous presence of efficiency in stone tool production for one raw material (quartzite) and inefficiency in stone tool production for another raw material (basalt) suggests that Oldowan producing hominins at DK were able to mediate the economic costs associated with stone tool procurement by utilizing high-cost materials more efficiently than is expected and low-cost materials in an inefficient manner.     

Vegetation and plant food reconstruction of lowermost Bed II, Olduvai Gorge, using modern analogs

Vegetation and plant foods for hominins of lowermost Bed II, Olduvai Gorge were modeled by examining vegetation in modern habitats in northern Tanzania (Lake Manyara, Ngorongoro, Serengeti) that are analogous to the paleolandscape in terms of climate, land forms, and soil types, as indicated by previous paleoenvironmental studies of Olduvai. Plant species in the modern habitats were identified in a series of sample plots, and those known to be eaten by modern humans, chimpanzees, or baboons were considered potentially edible for early hominins. Within the 50-80 kyr deposition of lowermost Bed II, periods of drier climate were characterized by low lake stands and a broad eastern lacustrine plain containing a mosaic of springs, marsh, woodland, and edaphic grassland. Based on results of this study, plant food diversity in each of those habitats was relatively low, but the mosaic nature of the area meant that hominins could reach several different habitat types within short distances, with access to potential plant foods including marsh plants, grass grains, roots, shrub fruits, edible parts from palms, leafy herbaceous plants, and Acacia pods, flowers, and gum. Based on Manyara analogs, a greater variety of plant foods, such as tree fruits (e.g., Ficus, Trichilia) and the roots and fruits of shrubs (e.g., Cordia, Salvadora) would be expected further east along the rivers in the lacustrine terrace and alluvial fans. Interfluves of the alluvial fans were probably less wooded and offered relatively fewer varieties of plant foods, but there is sparse paleoenvironmental evidence for the character of Olduvai's alluvial fans, making the choice of appropriate modern analogs difficult. In the western side of the basin, based on modern analogs in the Serengeti, riverine habitats provided the greatest variety of edible plant food species (e.g., Acacia, Grewia, Justicia). If the interfluves were grassland, then a large variety of potentially edible grasses and forbs were present seasonally. Periods of wetter climate resulted in a much expanded paleolake and a shrinking of the eastern lacustrine plain mosaic into a narrow zone. The dominant landscape features were then forest-lined rivers in the eastern alluvial fans, and rivers in the western side of the paleo-Olduvai basin were also better watered at these times, supporting denser woody vegetation with large varieties of edible fruits, leaves, and underground parts.     

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.     

A Homo habilis maxilla and other newly-discovered hominid fossils from Olduvai Gorge, Tanzania

In 1995, a 1.8 million year old hominid maxilla with complete dentition (OH 65) was excavated from Bed I in the western part of Olduvai Gorge. The molar crowns are small relative to the long flaring roots, and the root of the canine is very long and straight. The broad maxilla with wide U-shaped palate and the form of the tooth roots closely match those of KNM-ER 1470 which, in its parietal size and morphology, matches the type specimen of Homo habilis, OH 7. Thus, OH 65 and KNM-ER 1470 group with OH 7 as representatives of H. habilis while some other Olduvai specimens, such as OH 13 and OH 24, have more in common in terms of morphology and brain size with Australopithecus africanus. Between 1995 and 2007, the OLAPP team has recovered teeth of eight other hominid individuals from various parts of Olduvai Gorge. These have been identified as belonging to H. habilis, Paranthropus boisei, and Australopithecus cf. africanus.     

Unraveling hominin behavior at another anthropogenic site from Olduvai Gorge (Tanzania): new archaeological and taphonomic research at BK, Upper Bed II

New archaeological excavations and research at BK, Upper Bed II (Olduvai Gorge, Tanzania) have yielded a rich and unbiased collection of fossil bones. These new excavations show that BK is a stratified deposit formed in a riverine setting close to an alluvial plain. The present taphonomic study reveals the second-largest collection of hominin-modified bones from Olduvai, with abundant cut marks found on most of the anatomical areas preserved. Meat and marrow exploitation is reconstructed using the taphonomic signatures left on the bones by hominins. Highly cut-marked long limb shafts, especially those of upper limb bones, suggest that hominins at BK were actively engaged in acquiring small and middle-sized animals using strategies other than passive scavenging. The exploitation of large-sized game (Pelorovis) by Lower Pleistocene hominins, as suggested by previous researchers, is supported by the present study.     

Carbonate horizons, paleosols, and lake flooding cycles: Beds I and II of Olduvai Gorge, Tanzania

This study documents the petrology and stable isotope geochemistry of carbonates from six horizons from Beds I and II of Olduvai Gorge, Tanzania. The studied succession, immediately below and above Tuff IF, consists of interbedded waxy and earthy claystones with discrete carbonate horizons and thin sandstones. The succession was deposited in response to repeated flooding and withdrawal of a saline-alkaline lake. The carbonates and their overlying disconformities are important because they help define the surfaces on which hominin activity took place and allow very high-resolution correlation of geographically separated levels of hominin exploitation.The range of different carbonates includes unambiguous land-surface and pedogenic features including calcified rootmat horizons, rhizocretions, and micritic nodules, together with less determinate sparry calcite nodules. Stellate nodules are interpreted as pseudomorphs after sulfate-roses. The carbonate nodules are synsedimentary features, truncated by fluvial and other erosional surfaces. The isotopic composition of the carbonates is variable with δ¹⁸O ranging from −7.0‰ to −4.3‰, and δ¹³C from −8.5‰ to −1.6‰. A covariant increase in δ¹³C and δ¹⁸O repeats in each carbonate horizon and in individual nodules (inner to outer layers): it reflects the evolution of synsedimentary groundwaters. At times of low lake level, the carbonates started to precipitate from meteoric waters with low isotopic values and continued to form as lake levels rose and the waters became increasingly saline. Some of the samples have a last-stage cement of strontium rich dolomite, which supports late-stage flooding by the saline-alkaline lake. Previous studies of carbonate horizons from Olduvai have interpreted carbon isotope values in terms of changes in C₃ and C₄ plants that colonized the land surface. This study demonstrates that in some instances the isotope values from carbonates deposited in these lake marginal settings reflect changes in hydrology rather than vegetation.     

The O.H. 7 (Olduvai Gorge,Tanzania) hominid partial brain endocast revisited

The O.H.7 brain endocast was reexamined using a stereoplotting apparatus to quantify the amount of distortion. Some 127 measurements for each of the left and right sides were taken and treated statistically by student t-tests, both in paired and grouped fashion. Grouped data indicated no significant left-right differences. Paired data suggested three small regions of distortion, which produced three pairs of left-right differences that were significantly different. Given the very local nature of these minor distortions, it is concluded that the original reconstruction by Tobias was essentially correct. In addition, multiple regression analyses of selected chord-arc dimensions suggest that a volume in excess of 700 ml is most probable.     

First occurrence ofXerus cf.inauris (Rodentia,Sciruridae) at Olduvai Bed I (Lower Pleistocene,Tanzania)

PalZ - Some bonodont molars of a ground squirrel are attributed toXerus cf.inauris. Coming from Olduvai Bed I fossiliferous deposits they could represent an intermediate stage between the Laetoli...     

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.     

Examining time trends in the Oldowan technology at Beds I and II,Olduvai Gorge

The lithic analysis of the Bed I and II assemblages from Olduvai Gorge reveals both static and dynamic time trends in early hominids' technology from 1.8 to 1.2 m.y.a. The Bed I Oldowan (1.87-1.75 m.y.a.) is characterized by the least effort strategy in terms of raw material exploitation and tool production. The inclusion of new raw material, chert, for toolmaking in the following Developed Oldowan A (DOA, 1.65-1.53 m.y.a.) facilitated more distinctive and variable flaking strategies depending on the kind of raw materials. The unique characters of DOA are explainable by this raw material factor, rather than technological development of hominids. The disappearance of chert in the subsequent Developed Oldowan B and Acheulian (1.53-1.2 m.y.a.) necessitated a shift in tool production strategy more similar to that of Bed I Oldowan than DOA. However, the evidence suggests that Bed II hominids might have been more skillful toolmakers, intensive tool-users, and engaged in more active transport of stone tools than the Bed I predecessors. Koobi Fora hominids maintained a more static tool-using behavior than their Olduvai counterparts due mainly to a stable supply of raw materials. They differed from Olduvai hominids in terms of less battering of cores, consistent transport behavior, and few productions of side-struck flakes, indicating a regional variation of toolmaking and using practice. However, they shared with Olduvai hominids a temporal trend toward the production of larger flakes from larger cores after 1.6 m.y.a. Increased intake of animal resources and the expansion of ranging area of Homo ergaster would have led to the development of technological organization. Technological changes in the Oldowan industry are attested at Olduvai Gorge, Koobi Fora, and Sterkfontein, suggesting that it was a pan-African synchronous phenomenon, beginning at 1.5 m.y.a.