Age-depth model for uppermost Ndutu Beds constrains Middle Stone Age technology and climate-induced paleoenvironmental changes at Olduvai Gorge (Tanzania).

Olduvai Gorge in northern Tanzania is part of a globally important archeological and paleoanthropological World Heritage Site location critical to our understanding of modern human evolution. The Ndutu Beds in the upper part of the geological sequence at Olduvai Gorge represent the oldest unit to yield modern Homo sapiens skeletal material and Middle Stone Age technology. However, the timing of the deposition of the Ndutu Beds is poorly constrained at present, which limits our understanding of the paleoenvironments critical for contextualizing H. sapiens and related technologies in the Olduvai Basin. Using a suite of 15 luminescence ages of sedimentary core samples, combined with Bayesian statistics, this study provides a new higher-resolution age-depth model for the deposition of the uppermost Upper Ndutu and Naisiuiu Beds cored by the Olduvai Gorge Coring Project. The luminescence and modeled ages are presented as ±1 σ uncertainties. The Ndutu Beds intersected by the Olduvai Gorge Coring Project cores are dated to between 117.1 ± 17.9 and 45.3 ± 4.2 ka (between 125.9 ± 26.5 and 45.8 ± 8.2 ka modeled ages), while a probable overlying layer of Naisiusiu Beds dates to 23.7 ± 10.9 to 12.1 ± 1.7 ka (25.7 ± 18.9 ka and 12.0 ± 3.4 ka modeled age). Time-averaged accretion rates are derived during this time: (1) initially low rates (<5 cm ka-1) from the bottom of the core at 117.1 ± 17.9 ka up to 95.3 ± 11.1 ka (125.9 ± 26.5 to 95.5 ± 23.3 ka modeled ages); (2) the middle section spanning between 95.3 ± 11.1 and 62.7 ± 5.7 ka (95.5 ± 23.3 to 61.9 ± 10.4 ka modeled ages) with mean rates above 15 cm ka-1; and (3) the last 62.7 ± 5.7 ka (61.9 ± 10.4 ka modeled age) where the accretion rate reduces to below 5 cm ka-1. This reduction can be explained by the evolution of the gorge system that was likely driven by subsidence of the Olbalbal depression and changes in climate, particularly precipitation and resulting lake and base level changes. Older Upper Ndutu and Lower Ndutu Beds are contained within proto-gorges within the modern gorge system.

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.

The paleoecology of Pleistocene birds from Middle Bed II, at Olduvai Gorge, Tanzania, and the environmental context of the Oldowan-Acheulean transition.

Fossil bird data (community composition and taphonomic profiles) are used here to infer the environmental context of the Oldowan-Acheulean transitional period at Olduvai Gorge, Tanzania. This is the first comprehensive report on the Middle Bed II avifauna and includes fossils excavated by the Olduvai Geochronology and Archaeology Project (OGAP) and recently rediscovered fossils collected by Mary Leakey. Crane, ibis, darter, owl, raptor, crow, and vulture are reported from Bed II for the first time. The presence of these taxa, absent earlier in this Bed, point to a general opening and drying of the landscape with grassland and open woodland expansion. Taxa associated with dense, emergent wetland vegetation, such as dabbling ducks and rails, are uncommon and less diverse than earlier in Bed II. This suggests more mature wetlands with clearer waters. Cormorants continue to be common, but are less diverse. Cormorants and other roosting taxa provide evidence of trees in the area. Compared to lowermost Bed II, the Middle to Upper Bed II landscape is interpreted here as more open and drier (but not necessarily more arid), with matured wetlands, scattered trees, and a greater expansion of grasslands.

The carnivorous feeding behavior of early Homo at HWK EE, Bed II, Olduvai Gorge, Tanzania.

The regular consumption of large mammal carcasses, as evidenced by butchery marks on fossils recovered from Early Stone Age archaeological sites, roughly coincides with the appearance of Homo habilis. However, the significance of this niche expansion cannot be appreciated without an understanding of hominin feeding behavior and their ecological interactions with mammalian carnivores. The Olduvai Geochronology and Archaeology Project (OGAP) has recovered a large and well-preserved fossil assemblage from the HWK EE site, which was deposited just prior to the first appearance of Acheulean technology at Olduvai Gorge and likely represents one of the last H. habilis sites at Olduvai. This taphonomic analysis of the larger mammal fossil assemblage excavated from HWK EE shows evidence of multiple occupations over a long period of time, suggesting the site offered resources that were attractive to hominins. There was a water source indicated by the presence of fish, crocodiles, and hippos, and there was possible tree cover in an otherwise open habitat. The site preserves several stratigraphic intervals with large fossil and artifact assemblages within two of these intervals. Feeding traces on bone surfaces suggest hominins at the site obtained substantial amounts of flesh and marrow, particularly from smaller size group 1-2 carcasses, and exploited a wide range of taxa, including megafauna. A strong carnivore signal suggests hominins scavenged much of their animal foods during the two main stratigraphic intervals. In the later interval, lower carnivore tooth mark and hammerstone percussion mark frequencies, in addition to high epiphyseal to shaft fragment ratios, suggest hominins and carnivores did not fully exploit bone marrow and grease, which may have been acquired from nutritionally-stressed animals that died during a dry period at Olduvai. The diversity of fauna that preserve evidence of butchery suggests that the HWK EE hominins were opportunistic in their acquisition of carcass foods.

New excavations at the HWK EE site: Archaeology, paleoenvironment and site formation processes during late Oldowan times at Olduvai Gorge, Tanzania.

This paper reports the results of renewed fieldwork at the HWK EE site (Olduvai Gorge, Tanzania). HWK EE is positioned across the boundary between Lower and Middle Bed II, a crucial interval for studying the emergence of the Acheulean at Olduvai Gorge. Our excavations at HWK EE have produced one of the largest collections of fossils and artefacts from any Oldowan site, distributed across several archaeological units and a large excavation surface in four separate trenches that can be stratigraphically correlated. Here we present the main stratigraphic and archaeological units and discuss site formation processes. Results show a great density of fossils and stone tools vertically through two stratigraphic intervals (Lemuta and Lower Augitic Sandstone) and laterally across an area of around 300 m2, and highlight the confluence of biotic and abiotic agents in the formation of the assemblage. The large size and diversity of the assemblage, as well as its good preservation, qualify HWK EE as a reference site for the study of the late Oldowan at Olduvai Gorge and elsewhere in Africa. In addition, the description of the stratigraphic and archaeological sequence of HWK EE presented in this paper constitutes the foundation for further studies on hominin behavior and paleoecology in Lower and Middle Bed II.

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.

In situ ∼2.0 Ma trees discovered as fossil rooted stumps, lowermost Bed I, Olduvai Gorge, Tanzania.

The discovery of fossil rooted tree stumps in lowermost Lower Bed I from the western Olduvai Basin, Tanzania, age-bracketed by the Naabi Ignimbrite (2.038 ± 0.005 Ma) and Tuff IA (1.88 ± 0.05 Ma), provides the first direct, in situ, and to date oldest evidence of living trees at Olduvai Gorge. The tree relicts occur in an interval dominated by low-viscosity mass flow and braided fluvial sediments, deposited at the toe of a largely Ngorongoro Volcano-sourced volcaniclastic fan apron that comprised a widely spaced network of ephemeral braided streams draining northward into the Olduvai Basin. Preservation of the trees occurred through their engulfment by mass flows, post-mortem mold formation resulting from differential decay of woody tissues, and subsequent fluvially-related sediment infill, calcite precipitation, and cast formation. Rhizolith preservation was triggered by the interaction of root-induced organic and inorganic processes to form rhizocretionary calcareous root casts. Phytolith analyses were carried out to complete the paleoenvironmental reconstruction. They imply a pronounced seasonality and indicate a wooded landscape with grasses, shrubs, and sedges growing nearby, comparable to the low, open riverine woodland (unit 4c) along the Garusi River and tributaries in the Laetoli area. Among the tree stump cluster were found outsized lithic clasts and those consisting of quartzite were identified as Oldowan stone tool artifacts. In the context of hominin activity, the identification of wooded grassland in association with nearby freshwater drainages and Oldowan artifacts significantly extends our paleoenvironmental purview on the basal parts of Lower Bed I, and highlights the hitherto underrated role of the yet poorly explored western Olduvai Gorge area as a potential ecologically attractive setting and habitat for early hominins.

A one-million-year-old hominid distal ulna from Olduvai Gorge, Tanzania.

OBJECTIVE: Our aim was to recover new evidence of the evolution of the hominid lineage. METHODS: We undertook paleontological fieldwork at Olduvai Gorge, Tanzania, in one of the richest paleoanthropological sites in the world, documenting the evolution of our lineage and its environmental contexts over the last 2 million years. RESULTS: During field work in 2012, the Olduvai Vertebrate Paleontology Project discovered the distal end of a hominid ulna (OH 82) on the north side of Olduvai Gorge a few meters west of the Third Fault, eroding from Bed III sediments that are ∼1 million years in age. DISCUSSION: The size and morphology of this distal ulna falls within the normal range of variation seen in humans, although at the larger end of the distribution.

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.

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.

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.