Dietary options and behavior suggested by plant biomarker evidence in an early human habitat.

The availability of plants and freshwater shapes the diets and social behavior of chimpanzees, our closest living relative. However, limited evidence about the spatial relationships shared between ancestral human (hominin) remains, edible resources, refuge, and freshwater leaves the influence of local resources on our species' evolution open to debate. Exceptionally well-preserved organic geochemical fossils--biomarkers--preserved in a soil horizon resolve different plant communities at meter scales across a contiguous 25,000 m(2) archaeological land surface at Olduvai Gorge from about 2 Ma. Biomarkers reveal hominins had access to aquatic plants and protective woods in a patchwork landscape, which included a spring-fed wetland near a woodland that both were surrounded by open grassland. Numerous cut-marked animal bones are located within the wooded area, and within meters of wetland vegetation delineated by biomarkers for ferns and sedges. Taken together, plant biomarkers, clustered bone debris, and hominin remains define a clear spatial pattern that places animal butchery amid the refuge of an isolated forest patch and near freshwater with diverse edible resources.

Ecosystem variability and early human habitats in eastern Africa.

The role of savannas during the course of early human evolution has been debated for nearly a century, in part because of difficulties in characterizing local ecosystems from fossil and sediment records. Here, we present high-resolution lipid biomarker and isotopic signatures for organic matter preserved in lake sediments at Olduvai Gorge during a key juncture in human evolution about 2.0 Ma--the emergence and dispersal of Homo erectus (sensu lato). Using published data for modern plants and soils, we construct a framework for ecological interpretations of stable carbon-isotope compositions (expressed as δ(13)C values) of lipid biomarkers from ancient plants. Within this framework, δ(13)C values for sedimentary leaf lipids and total organic carbon from Olduvai Gorge indicate recurrent ecosystem variations, where open C(4) grasslands abruptly transitioned to closed C(3) forests within several hundreds to thousands of years. Carbon-isotopic signatures correlate most strongly with Earth's orbital geometry (precession), and tropical sea-surface temperatures are significant secondary predictors in partial regression analyses. The scale and pace of repeated ecosystem variations at Olduvai Gorge contrast with long-held views of directional or stepwise aridification and grassland expansion in eastern Africa during the early Pleistocene and provide a local perspective on environmental hypotheses of human evolution.

Water, plants, and early human habitats in eastern Africa.

Water and its influence on plants likely exerted strong adaptive pressures in human evolution. Understanding relationships among water, plants, and early humans is limited both by incomplete terrestrial records of environmental change and by indirect proxy data for water availability. Here we present a continuous record of stable hydrogen-isotope compositions (expressed as δD values) for lipid biomarkers preserved in lake sediments from an early Pleistocene archaeological site in eastern Africa--Olduvai Gorge. We convert sedimentary leaf- and algal-lipid δD values into estimates for ancient source-water δD values by accounting for biochemical, physiological, and environmental influences on isotopic fractionation via published water-lipid enrichment factors for living plants, algae, and recent sediments. Reconstructed precipitation and lake-water δD values, respectively, are consistent with modern isotopic hydrology and reveal that dramatic fluctuations in water availability accompanied ecosystem changes. Drier conditions, indicated by less negative δD values, occur in association with stable carbon-isotopic evidence for open, C(4)-dominated grassland ecosystems. Wetter conditions, indicated by lower δD values, are associated with expanded woody cover across the ancient landscape. Estimates for ancient precipitation amounts, based on reconstructed precipitation δD values, range between approximately 250 and 700 mm · y(-1) and are consistent with modern precipitation data for eastern Africa. We conclude that freshwater availability exerted a substantial influence on eastern African ecosystems and, by extension, was central to early human proliferation during periods of rapid climate change.

A spring forward for hominin evolution in East Africa.

Groundwater is essential to modern human survival during drought periods. There is also growing geological evidence of springs associated with stone tools and hominin fossils in the East African Rift System (EARS) during a critical period for hominin evolution (from 1.8 Ma). However it is not known how vulnerable these springs may have been to climate variability and whether groundwater availability may have played a part in human evolution. Recent interdisciplinary research at Olduvai Gorge, Tanzania, has documented climate fluctuations attributable to astronomic forcing and the presence of paleosprings directly associated with archaeological sites. Using palaeogeological reconstruction and groundwater modelling of the Olduvai Gorge paleo-catchment, we show how spring discharge was likely linked to East African climate variability of annual to Milankovitch cycle timescales. Under decadal to centennial timescales, spring flow would have been relatively invariant providing good water resource resilience through long droughts. For multi-millennial periods, modelled spring flows lag groundwater recharge by 100 s to 1000 years. The lag creates long buffer periods allowing hominins to adapt to new habitats as potable surface water from rivers or lakes became increasingly scarce. Localised groundwater systems are likely to have been widespread within the EARS providing refugia and intense competition during dry periods, thus being an important factor in natural selection and evolution, as well as a vital resource during hominin dispersal within and out of Africa.

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.

Stable isotopes of pedogenic carbonates as indicators of paleoecology in the Plio-Pleistocene (upper Bed I), western margin of the Olduvai Basin, Tanzania.

Paleosol carbonates from trenches excavated as part of a landscape-scale project in Bed I of Olduvai Gorge, Tanzania, were analyzed for stable carbon and oxygen isotopic composition. The approximately 60,000-year interval ( approximately 1.845-1.785 Ma) above Tuff IB records evidence for lake and fluvial sequences, volcanic eruptions, eolian and pedogenic processes, and the development of a fluvial plain in the western margin of the basin. Significant temporal variation in the carbonate delta(18)O values records variation of local precipitation and supports the shifts in climatic conditions interpreted from the lithologic record. During this period, carbonate delta(13)C values varied between depositional facies indicating that the paleolandscape supported a local biomass of about 40-60% C(4) plants within a mosaic of grassy woodlands and wooded grasslands. The lithologic and stable isotope record in this small lake basin indicates the area was much wetter, with more woody C(3) plants, during this interval than is the semi-arid area today. The record also reflects the variation in climatic conditions (wet/dry) documented by other global climate proxies for this time.

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.