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.     

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.     

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.     

Cranial remains of Homo erectus from Beds II and IV,Olduvai Gorge,Tanzania

Cranial remains of hominids 9 and 12 from Olduvai Gorge are described in detail. O.H. 9 consists of a heavily built braincase, partly damaged and lacking the face, while O.H. 12 is less complete. The Bed II specimen is about 1.2 million years in age and shows anatomical similarities to the cranium designated ER-3733 from Koobi Fora, east of Lake Turkana. Together these African fossils provide valuable information about Homo erectus in the later Lower Pleistocene. Comparisons of O.H. 9 with several of the Choukoutien crania are also carried out. These Chinese and other Asian remains of Homo erectus cannot be placed in a secure chronological framework, but all of the material should be studied systematically in order to assess relatedness among what must be several different populations.     

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.     

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.     

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.     

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.     

First Partial Skeleton of a 1.34-Million-Year-Old Paranthropus boisei from Bed II,Olduvai Gorge,Tanzania

Recent excavations in Level 4 at BK (Bed II, Olduvai Gorge, Tanzania) have yielded nine hominin teeth, a distal humerus fragment, a proximal radius with much of its shaft, a femur shaft, and a tibia shaft fragment (cataloged collectively as OH 80). Those elements identified more specifically than to simply Hominidae gen. et sp. indet are attributed to Paranthropus boisei. Before this study, incontrovertible P. boisei partial skeletons, for which postcranial remains occurred in association with taxonomically diagnostic craniodental remains, were unknown. Thus, OH 80 stands as the first unambiguous, dentally associated Paranthropus partial skeleton from East Africa. The morphology and size of its constituent parts suggest that the fossils derived from an extremely robust individual who, at 1.338±0.024 Ma (1 sigma), represents one of the most recent occurrences of Paranthropus before its extinction in East Africa.     

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.     

A new species of cormorant (Aves: Phalacrocoracidae) from the Pleistocene of Olduvai Gorge,Tanzania

Phalacrocorax owrei nov. sp. is a small cormorant from the Lower Pleistocene of Olduvai Gorge. Osteological proportions are established for the four subgenera of Phalacrocorax, and P. owrei is assigned to the subgenus Stictocarbo. The species was the most abundant bird in the Bed I deposits and is also represented by a few specimens in the middle part of Bed II. Its last known appearance coincides with a change in the local environment when the climate became more arid and the Olduvai Lake became more saline and more alkaline. At other localities in Bed II the extinct P. tanzanice occurs, as well as P. africanus and P. corbo, which breed on the African lakes and seacoast today.     

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.     

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.     

Fingerprinting facies of the Tuff IF marker,with implications for early hominin palaeoecology,Olduvai Gorge,Tanzania

The top of Tuff IF marks the upper boundary of the Plio-/Pleistocene Bed I succession exposed in Olduvai Gorge, NE Tanzania, a tephrostratigraphic interval that is well known for remarkable Oldowan archaeological and vertebrate fossil assemblages, including early hominins. Geochemically, Tuff IF is characterized by a relatively consistent silica-undersaturated trachytic to phonolitic composition that relates both in terms of numerical ages (1.79 Ma) and compositional constraints to Olmoti volcano, located 22–38 km E of Olduvai Gorge.Measured Tuff IF sections are characterized by a thick unwelded pyroclastic flow in proximal settings and a succession of surges and ashfalls that interfingers with fluvio-lacustrine deposits of the Olduvai Basin in the medial to distal settings. Only in those areas with a relatively low topographic gradient, in distal reaches and beyond the toe of an Olmoti-sourced volcaniclastic fan, did the Tuff IF marker develop a typical threefold subdivision comprising: (1) primary surges and minor fallout, (2) reworked pumice units as lateral correlatives of proximally emplaced pyroclastic flows and (3) a succession of mass flows in conjunction with aeolian and fluvially reworked units.The pyroclastic marker unit is thus highly heterogeneous with regard to its vertical and lateral facies architecture and this has immediate effects on its preservation potential and in situ burial of fossils and stone artifacts. Even though the volcanically-related environmental perturbations were probably more severe at the eastern lake margin, the evidence for at least temporary freshwater sources and trees suggests environments conducive to hominin activities, but not during emplacement of Tuff IF pyroclastic flows and surges. This inference is supported by the presence of rich Oldowan stone artifact assemblages immediately preceding and following the deposition of Tuff IF, but only extremely sparse archaeological traces from one site in Tuff IF, restricted to the upper, fluvially reworked portion of Tuff IF.Tuff IF facies record the maximum of a regressive (drying) cycle that correlates with regional marine arid indicators. An ecological crisis covering ～ 2000–3500 a of time during Tuff IF deposition resulted not only from the lethal effects of explosive volcanism but also from its coincidence with a pronounced period of climate induced drought. These combined effects appear to have made at least the eastern basin uninhabitable by hominins and other vertebrates for most of the time during which Tuff IF accumulated.     

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.     

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.