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Understanding the climatic drivers of environmental variability (EV) during the Plio-Pleistocene and EV’s influence on mammalian macroevolution are two outstanding foci of research in African paleoclimatology and evolutionary biology. The potential effects of EV are especially relevant for testing the variability selection hypothesis, which predicts a positive relationship between EV and speciation and extinction rates in fossil mammals. Addressing these questions is stymied, however, by 1) a lack of multiple comparable EV records of sufficient temporal resolution and duration, and 2) the incompleteness of the mammalian fossil record. Here, we first compile a composite history of Pan-African EV spanning the Plio-Pleistocene, which allows us to explore which climatic variables influenced EV. We find that EV exhibits 1) a long-term trend of increasing variability since ∼3.7 Ma, coincident with rising variability in global ice volume and sea surface temperatures around Africa, and 2) a 400-ky frequency correlated with seasonal insolation variability. We then estimate speciation and extinction rates for fossil mammals from eastern Africa using a method that accounts for sampling variation. We find no statistically significant relationship between EV and estimated speciation or extinction rates across multiple spatial scales. These findings are inconsistent with the variability selection hypothesis as applied to macroevolutionary processes.

Understanding eastern African paleoclimate is critical for contextualizing early human evolution, adaptation, and dispersal, yet Pleistocene climate of this region and its governing mechanisms remain poorly understood due to the lack of long, orbitally-resolved, terrestrial paleoclimate records. Here we present leaf wax hydrogen isotope records of rainfall from paleolake sediment cores from key time windows that resolve long-term trends, variations, and high-latitude effects on tropical African precipitation. Eastern African rainfall was dominantly controlled by variations in low-latitude summer insolation during most of the early and middle Pleistocene, with little evidence that glacial-interglacial cycles impacted rainfall until the late Pleistocene. We observe the influence of high-latitude-driven climate processes emerging from the last interglacial (Marine Isotope Stage 5) to the present, an interval when glacial-interglacial cycles were strong and insolation forcing was weak. Our results demonstrate a variable response of eastern African rainfall to low-latitude insolation forcing and high-latitude-driven climate change, likely related to the relative strengths of these forcings through time and a threshold in monsoon sensitivity. We observe little difference in mean rainfall between the early, middle, and late Pleistocene, which suggests that orbitally-driven climate variations likely played a more significant role than gradual change in the relationship between early humans and their environment., (© 2022. The Author(s).)

Hominin fire use in the early Pleistocene has been debated since the early 1970s when consolidated reddened sediment patches were identified at FxJj20 East and Main, Koobi Fora, Kenya. Since then, researchers have argued for evidence of early Pleistocene fire use at a handful of archaeological sites with evidence of combustion. Some argue that morphological evidence of early Homo erectus fossils indicates a dietary shift to higher quality food sources, which could be achieved by cooking. Others contend that fire use does not become a regular behavior until later, in the middle Pleistocene, when archaeological sites begin to show regular evidence for fire use. An early date for hominin control of fire would help to explain the grade changes seen with the appearance of H. erectus, while a later date would mean that fire would have had little influence on the early development of the lineage. Early hominins would have encountered fire regularly on the landscape, increasing the possibility of hominins interacting with and habituating to natural landscape fire. Only a detailed understanding of the patterns of controlled and natural fires can lead to understanding of early hominin fire use. We present new work on the evidence of fire at the FxJj20 Site complex in Koobi Fora, dated to 1.5 Ma. We highlight evidence of burning found on site through Fourier Transform Infrared spectrometry, and describe ongoing work to investigate the association of hominin behavior and fire evidence. We present data supporting the hypothesis that the site is undisturbed and discuss spatial relationships showing burned material associated with non-burned material. We present data on a type of stone fragment, the Thermal Curve Fragment (TCF), which is indicative of knapped material being exposed to high heat. Finally, we suggest future directions on the topic of fire in the early Pleistocene., (Copyright © 2019 Elsevier Ltd. All rights reserved.)

The influence of climate change on hominin evolution is much debated. Two issues hamper our understanding of this process: the limited hominin fossil record, and incomplete knowledge about hominin spatial occupation of Africa. Here, we analyze the presently known hominin fossil distribution pattern and explore the potential geographic distribution of hominins between ∼4.5 and ∼2.5 Ma. We focus on assessing the relevance of the Coastal Forest of Eastern Africa (CFEA) along the Indian Ocean as a core area for early hominin evolution. Based on biogeographic-phylogeographic data we propose the coastal refuge hypothesis: the CFEA provided a refugium for early hominins in periods of variable climate and strong seasonality during eccentricity maxima. From this refuge, evolved species could disperse inland (e.g. to rift basins) via vegetated humid corridors, whenever onset of stable climate periods with low seasonality during eccentricity minima allowed expansion out of the coastal enclave. We develop a conceptual model in time and space, comparing predictions with climatic and hominin fossil records. The results imply that: 1) between ∼4.5 and 3 Ma, ongoing (mostly anagenetic) hominin evolution occurred in the CFEA, punctuated by inland dispersal events at ∼4.4, 4.2, 3.8, 3.5, and 3.2 Ma; 2) before ∼3 Ma, the Afar Basin was a (sub)core area often connected to and relatively similar to the CFEA, while other inland areas were more or less marginal for early hominin habitation; 3) after ∼3 Ma, Northern Hemisphere Glaciation exerted strong influence by causing latitudinal contraction of the CFEA, leading to habitat fragmentation, isolation of hominin populations and possible cladogenetic evolution. A major challenge for the coastal refuge model is the fact that at present, no (hominin) fossils are known from the CFEA. We consider how this can be explained, and possibly overcome with targeted search efforts. Furthermore we discuss how the model can be tested, e.g. with molecular phylogeography approaches, and used to predict new hominin fossil locations. With this study, we hope to contribute a fresh perspective to the climate-evolution debate, emphasizing the role of climatic stability, length of dry season and vegetation cover to facilitate connectivity between hominin core and marginal habitats., (Copyright © 2019 The Authors. Published by Elsevier Ltd.. All rights reserved.)

Human evolutionary scholars have long supposed that the earliest stone tools were made by the genus Homo and that this technological development was directly linked to climate change and the spread of savannah grasslands. New fieldwork in West Turkana, Kenya, has identified evidence of much earlier hominin technological behaviour. We report the discovery of Lomekwi 3, a 3.3-million-year-old archaeological site where in situ stone artefacts occur in spatiotemporal association with Pliocene hominin fossils in a wooded palaeoenvironment. The Lomekwi 3 knappers, with a developing understanding of stone's fracture properties, combined core reduction with battering activities. Given the implications of the Lomekwi 3 assemblage for models aiming to converge environmental change, hominin evolution and technological origins, we propose for it the name 'Lomekwian', which predates the Oldowan by 700,000 years and marks a new beginning to the known archaeological record.

During the evolution of hominins, it is generally accepted that there was a shift in postcranial morphology between Australopithecus and the genus Homo. Given the scarcity of associated remains of early Homo, however, relatively little is known about early Homo postcranial morphology. There are hints of postcranial diversity among species, but our knowledge of the nature and extent of potential differences is limited. Here we present a new associated partial ilium and femur from Koobi Fora, Kenya, dating to 1.9 Ma (millions of years ago) that is clearly attributable to the genus Homo but documents a pattern of morphology not seen in eastern African early Homo erectus. The ilium and proximal femur share distinctive anatomy found only in Homo. However, the geometry of the femoral midshaft and contour of the pelvic inlet do not resemble that of any specimens attributed to H. erectus from eastern Africa. This new fossil confirms the presence of at least two postcranial morphotypes within early Homo, and documents diversity in postcranial morphology among early Homo species that may reflect underlying body form and/or adaptive differences., (Copyright © 2015 Elsevier Ltd. All rights reserved.)

To address questions regarding the evolutionary origin, radiation and dispersal of the genus Homo, it is crucial to be able to place the occurrence of hominin fossils in a high-resolution chronological framework. The period around 2 Ma (millions of years ago) in eastern Africa is of particular interest as it is at this time that a more substantial fossil record of the genus Homo is first found. Here we combine magnetostratigraphy and strontium (Sr) isotope stratigraphy to improve age control on hominin-bearing upper Burgi (UBU) deposits in Areas 105 and 131 on the Karari Ridge in the eastern Turkana Basin (Kenya). We identify the base of the Olduvai subchron (bC2n) plus a short isolated interval of consistently normal polarity that we interpret to be the Pre-Olduvai event. Combined with precession-forced (~20 kyr [thousands of years]) wet-dry climate cycles resolved by Sr isotope ratios, the magnetostratigraphic data allow us to construct an age model for the UBU deposits. We provide detailed age constraints for 15 hominin fossils from Area 131, showing that key specimens such as cranium KNM-ER 1470, partial face KNM-ER 62000 and mandibles KNM-ER 1482, KNM-ER 1801, and KNM-ER 1802 can be constrained between 1.945 ± 0.004 and 2.058 ± 0.034 Ma, and thus older than previously estimated. The new ages are consistent with a temporal overlap of two species of early Homo that can be distinguished by their facial morphology. Further, our results show that in this time interval, hominins occurred throughout the wet-dry climate cycles, supporting the hypothesis that the lacustrine Turkana Basin was a refugium during regionally dry periods. By establishing the observed first appearance datum of a marine-derived stingray in UBU deposits at 2.058 ± 0.034 Ma, we show that at this time the Turkana Basin was hydrographically connected to the Indian Ocean, facilitating dispersal of fauna between these areas. From a biogeographical perspective, we propose that the Indian Ocean coastal strip should be considered as a possible source area for one or more of the multiple Homo species in the Turkana Basin from over 2 Ma onwards., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

The origin and evolution of early Pleistocene hominin lithic technologies in Africa occurred within the context of savanna grassland ecosystems. The Nachukui Formation of the Turkana Basin in northern Kenya, containing Oldowan and Acheulean tool assemblages and fossil evidence for early members of Homo and Paranthropus, provides an extensive spatial and temporal paleosol record of early Pleistocene savanna flora. Here we present new carbon isotopic (δ(13)CVPDB) values of pedogenic carbonates (68 nodules, 193 analyses) from the Nachukui Formation in order to characterize past vegetation structure and change through time. We compared three members (Kalochoro, Kaitio, and Natoo) at five locations spanning 2.4-1.4Ma and sampled in proximity to hominin archaeological and paleontological sites. Our results indicate diverse habitats showing a mosaic pattern of vegetation cover at each location yet demonstrate grassland expansion through time influenced by paleogeography. Kalochoro floodplains occurred adjacent to large river systems, and paleosols show evidence of C3 woodlands averaging 46-50% woody cover. Kaitio habitats were located along smaller rivers and lake margins. Paleosols yielded evidence for reduced portions of woody vegetation averaging 34-37% woody cover. Natoo environments had the highest percentage of grasslands averaging 21% woody cover near a diminishing Lake Turkana precursor. We also compared paleosol δ(13)CVPDB values of lithic archaeological sites with paleosol δ(13)CVPDB values of all environments available to hominins at 2.4-1.4Ma in the Nachukui and Koobi Fora Formations. Grassy environments became more widespread during this interval; woody canopy cover mean percentages steadily decreased by 12%. However, significantly more wooded savanna habitats were present in the vicinity of lithic archaeological sites and did not mirror the basin-wide trend of grassland spread. Hominin lithic archaeological sites consistently demonstrated woody cover circa 40% throughout our study interval and were 4-12% more woody than coeval basin environs. We propose that Turkana Basin early tool makers may have preferred a more wooded portion of the savanna ecosystem to reduce heat stress and to gain differential access to potable water, raw materials, animal carcasses, and edible plants., (Copyright © 2013 Elsevier Ltd. All rights reserved.)

Since its discovery in 1972 (ref. 1), the cranium KNM-ER 1470 has been at the centre of the debate over the number of species of early Homo present in the early Pleistocene epoch of eastern Africa. KNM-ER 1470 stands out among other specimens attributed to early Homo because of its larger size, and its flat and subnasally orthognathic face with anteriorly placed maxillary zygomatic roots. This singular morphology and the incomplete preservation of the fossil have led to different views as to whether KNM-ER 1470 can be accommodated within a single species of early Homo that is highly variable because of sexual, geographical and temporal factors, or whether it provides evidence of species diversity marked by differences in cranial size and facial or masticatory adaptation. Here we report on three newly discovered fossils, aged between 1.78 and 1.95 million years (Myr) old, that clarify the anatomy and taxonomic status of KNM-ER 1470. KNM-ER 62000, a well-preserved face of a late juvenile hominin, closely resembles KNM-ER 1470 but is notably smaller. It preserves previously unknown morphology, including moderately sized, mesiodistally long postcanine teeth. The nearly complete mandible KNM-ER 60000 and mandibular fragment KNM-ER 62003 have a dental arcade that is short anteroposteriorly and flat across the front, with small incisors; these features are consistent with the arcade morphology of KNM-ER 1470 and KNM-ER 62000. The new fossils confirm the presence of two contemporary species of early Homo, in addition to Homo erectus, in the early Pleistocene of eastern Africa.

The Turkana Basin preserves a long and detailed record of biotic evolution, cultural development, and rift valley geology in its sedimentary strata. Before the formation of the modern basin, Cretaceous fluvial systems, Paleogene lakes, and Oligo-Miocene volcano-sedimentary sequences left fossil-bearing strata in the region. These deposits were in part related to an early system of rift basins that stretched from Sudan to the Indian Ocean. The present-day basin has its origins in Pliocene tectonic developments of the modern rift, with subsidence making room for more than one kilometer of Plio-Pleistocene strata. Much of this sequence belongs to the Omo Group, richly fossiliferous sediments associated with the ancestral Omo River and its tributaries. Modern Lake Turkana has a record stretching back more than 200 thousand years, with earlier lake phases throughout the Plio-Pleistocene. The geologic history of the basin is one of dynamic landscapes responding to environmental influences, including tectonics, volcanic activity and climate., (Copyright © 2011 Wiley Periodicals, Inc.)

The Acheulian is one of the first defined prehistoric techno-complexes and is characterized by shaped bifacial stone tools. It probably originated in Africa, spreading to Europe and Asia perhaps as early as ∼1 million years (Myr) ago. The origin of the Acheulian is thought to have closely coincided with major changes in human brain evolution, allowing for further technological developments. Nonetheless, the emergence of the Acheulian remains unclear because well-dated sites older than 1.4 Myr ago are scarce. Here we report on the lithic assemblage and geological context for the Kokiselei 4 archaeological site from the Nachukui formation (West Turkana, Kenya) that bears characteristic early Acheulian tools and pushes the first appearance datum for this stone-age technology back to 1.76 Myr ago. Moreover, co-occurrence of Oldowan and Acheulian artefacts at the Kokiselei site complex indicates that the two technologies are not mutually exclusive time-successive components of an evolving cultural lineage, and suggests that the Acheulian was either imported from another location yet to be identified or originated from Oldowan hominins at this vicinity. In either case, the Acheulian did not accompany the first human dispersal from Africa despite being available at the time. This may indicate that multiple groups of hominins distinguished by separate stone-tool-making behaviours and dispersal strategies coexisted in Africa at 1.76 Myr ago.

Homo erectus was the first human lineage to disperse widely throughout the Old World, the only hominin in Asia through much of the Pleistocene, and was likely ancestral to H. sapiens. The demise of this taxon remains obscure because of uncertainties regarding the geological age of its youngest populations. In 1996, some of us co-published electron spin resonance (ESR) and uranium series (U-series) results indicating an age as young as 35-50 ka for the late H. erectus sites of Ngandong and Sambungmacan and the faunal site of Jigar (Indonesia). If correct, these ages favor an African origin for recent humans who would overlap with H. erectus in time and space. Here, we report (40)Ar/(39)Ar incremental heating analyses and new ESR/U-series age estimates from the "20 m terrace" at Ngandong and Jigar. Both data sets are internally consistent and provide no evidence for reworking, yet they are inconsistent with one another. The (40)Ar/(39)Ar analyses give an average age of 546±12 ka (sd±5 se) for both sites, the first reliable radiometric indications of a middle Pleistocene component for the terrace. Given the technical accuracy and consistency of the analyses, the argon ages represent either the actual age or the maximum age for the terrace and are significantly older than previous estimates. Most of the ESR/U-series results are older as well, but the oldest that meets all modeling criteria is 143 ka+20/-17. Most samples indicated leaching of uranium and likely represent either the actual or the minimum age of the terrace. Given known sources of error, the U-series results could be consistent with a middle Pleistocene age. However, the ESR and (40)Ar/(39)Ar ages preclude one another. Regardless, the age of the sites and hominins is at least bracketed between these estimates and is older than currently accepted.

Geological data from the Bura Hasuma region at Koobi Fora provide important constraints for estimating the ages of hominin fossils recovered there, including the cranium KNM-ER 1813. Strata of the upper Burgi, KBS, and Okote members in this part of Koobi Fora reflect three depositional regimes driven by changing paleogeography through time. The upper Burgi and lowermost KBS sequence in the southern Bura Hasuma region accumulated in a lacustrine to delta front setting, with highly localized depositional patterns, limiting the lateral extent of lithostratigraphic markers. Farther north, uppermost upper Burgi through KBS member strata document a fluctuating lake margin, with complex facies patterns. This interval is marked by laterally extensive lithostratigraphic markers, including molluscan packstones, beach sandstones, and stromatolite beds. The uppermost KBS and Okote members show a transition to dominantly fluvial character, with localized and discontinuous accumulation. An age model for the richly fossiliferous Area 123 sequence demonstrates the complexity of terrestrial accumulation patterns. Early lacustrine and delta front accumulation is marked by fairly continuous sedimentation, and high accumulation rates (up to ca. 91 cm/k.yr.). The fluctuating lake margin interval reflects lower sedimentation rates coupled with intervals of exposure, decreasing accumulation significantly (to ca. 13 cm/k.yr.). The capping fluvial interval is marked by significant erosion surfaces, breaks which may drop the overall accumulation rate even lower (ca. 0.3 cm/k.yr.). The data provided here establish a geological framework at odds with a recent proposal of ages considerably younger (by ca. 250 k.yr.) for many of the fossils from Area 123 and elsewhere. Tests of age models demonstrate that the younger ages are not possible. While minor refinements to age estimates for fossils are indicated by improved chronostratigraphic control, in the case of KNM-ER 1813, an age of younger than 1.78 Ma is precluded on magnetostratigraphic grounds.

Detailed stratigraphic analysis of the Omo I and Omo II fossil localities confirms both the relational and sequential context reported by Butzer in 1969. The two fossils derive from approximately the same level within upper Member I of the Kibish Formation. Additional features of the local stratigraphic sequences indicate a complex history of depositional events, minor erosional surfaces, and weak soil formation throughout upper Member I.

A running controversy in evolutionary thought was Eldredge and Gould's punctuated equilibrium model, which proposes long periods of morphological stasis interspersed with rapid bursts of dramatic evolutionary change. One of the earliest and most iconic pieces of research in support of punctuated equilibrium is the work of Williamson on the Plio-Pleistocene molluscs of the Turkana Basin. Williamson claimed to have found firm evidence for three episodes of rapid evolutionary change separated by long periods of stasis in a high-resolution sequence. Most of the discussions following this report centered on the topics of (eco)phenotype versus genotype and the possible presence of preservational and temporal artifacts. The debate proved inconclusive, leaving Williamson's reports as one of the empirical foundations of the paradigm of punctuated equilibrium. Here we conclusively show Williamson's original interpretations to be highly flawed. The supposed rapid bursts of punctuated evolutionary change represent artifacts resulting from the invasion of extrabasinal faunal elements in the Turkana palaeolakes during wet phases well known from elsewhere in Africa.

Central to the debate surrounding global climate change and Plio-Pleistocene hominin evolution is the degree to which orbital-scale climate patterns influence low-latitude continental ecosystems and how these influences can be distinguished from regional volcano-tectonic events and local environmental effects. The Pliocene Hadar Formation of Ethiopia preserves a record of hominin paleoenvironments from roughly 3.5 to 2.2 Ma at a temporal resolution relevant to evolutionary change within hominins and other taxa. This study integrates the high-resolution sedimentological and paleontological records at Hadar with climate proxies such as marine core isotope, dust, and sapropel records. Consistent cycling observed both between and within fluvial and lacustrine depositional environments prior to 2.9 Ma at Hadar appears to be predominantly climatic in nature. In contrast a significant change in depositional facies after 2.9 Ma to sequences dominated by conglomerate cut-and-fill cycles indicates a strong tectonic signature related to regional developments in the Main Ethiopian Rift. While specific events seen in marine proxy records may have parallels in the Hadar environmental archive, their overall patterns of high versus low variability may be even more relevant. For example, periods of relatively high-amplitude climate oscillations between 3.15 and 2.95 Ma may be linked to noted size-related morphological changes within the Hadar Australopithecus afarensis lineage and a significant increase in more arid-adapted bovid taxa. Increased aridity in East Africa during this period is also indicated by peaks in eolian dust in the marine core record. Conversely, the dominant lacustrine phase at Hadar ca. 3.3 Ma coincides with the least variable period in several climate proxy records, including marine core foraminifera delta(18)O values and eolian dust concentration. This phase is also coeval with low insolation variability and a very distinct and significant long-term period of low dust percentage in circum-Africa marine cores.

Plio-Pleistocene East African grassland expansion and faunal macroevolution, including that of our own lineage, are attributed to global climate change. To further understand environmental factors of early hominin evolution, we reconstruct the paleogeographic distribution of vegetation (C(3)-C(4) pathways) by stable carbon isotope (delta(13)C) analysis of pedogenic carbonates from the Plio-Pleistocene Koobi Fora region, northeast Lake Turkana Basin, Kenya. We analyzed 202 nodules (530 measurements) from ten paleontological/archaeological collecting areas spanning environments over a 50-km(2) area. We compared results across subregions in evolving fluviolacustrine depositional environments in the Koobi Fora Formation from 2.0-1.5 Ma, a stratigraphic interval that temporally brackets grassland ascendancy in East Africa. Significant differences in delta(13)C values between subregions are explained by paleogeographic controls on floral composition and distribution. Our results indicate grassland expansion between 2.0 and 1.75 Ma, coincident with major shifts in basin-wide sedimentation and hydrology. Hypotheses may be correct in linking Plio-Pleistocene hominin evolution to environmental changes from global climate; however, based on our results, we interpret complexity from proximate forces that mitigated basin evolution. An approximately 2.5 Ma tectonic event in southern Ethiopia and northern Kenya exerted strong effects on paleography in the Turkana Basin from 2.0-1.5 Ma, contributing to the shift from a closed, lacustrine basin to one dominated by open, fluvial conditions. We propose basin transformation decreased residence time for Omo River water and expanded subaerial floodplain landscapes, ultimately leading to reduced proportions of wooded floras and the establishment of habitats suitable for grassland communities.

Climate change is hypothesized as a cause of major events of Plio-Pleistocene East African hominin evolution, but the vertically discontinuous and laterally confined nature of the relevant geological records has led to difficulties with assessing probable links between the two. High-resolution sedimentary sequences from lacustrine settings can provide comprehensive data of environmental changes and detailed correlations with well-established orbital and marine records of climate. Hominin-bearing deposits from Koobi Fora Ridge localities in the northeast Turkana Basin of Kenya are an archive of Plio-Pleistocene lake-margin sedimentation though significant developmental junctures of northern African climates, East African environments, and hominin evolution. This study examines alluvial channel and floodplain, nearshore lacustrine, and offshore lacustrine facies environments for the approximately 136-m-thick KBS Member (Koobi Fora Formation) exposed at the Koobi Fora Ridge. Aspects of the facies environments record information on the changing hydrosedimentary dynamics of the lake margin and give insights into potential climatic controls. Seasonal/yearly climate changes are represented by the varve-like laminations in offshore mudstones and the slickensides, dish-shaped fractures, and other paleosol features overprinted on floodplain strata. Vertical shifts between facies environments, however, are interpreted to indicate lake-level fluctuations deriving from longer-term, dry-wet periods in monsoonal rainfall. Recurrence periods for the inferred lake-level changes range from about 10,000 to 50,000 years, and several are consistent with the average estimated timescales of orbital precession ( approximately 20,000 years) and obliquity ( approximately 40,000 years). KBS Member facies environments from the Koobi Fora Ridge document the development of lake-margin hominin habitats in the northeast Turkana Basin. Environmental changes in these habitats may be a result of monsoonal rainfall variations that derive from orbital insolation and/or glacial forcing.

The Acheulean site of Gesher Benot Ya'aqov in the Dead Sea Rift of Israel documents hominin movements and technological development on a corridor between Africa and Eurasia. New age data place the site at 780,000 years ago (oxygen isotope stage 19), considerably older than previous estimates. The archaeological data from the site portray strong affinities with African stone tool traditions. The findings also reflect adroit technical skills and in-depth planning abilities, more advanced and complex than those of earlier archaeological occurrences in the Levant.

The discovery of Australopithecus anamensis fossils from strata lying between tephra dated at 4.17 and 4.12 million years ago, and from slightly higher strata not well constrained in age by overlying dated units, provoked the claim that more than one species might be represented: it was suggested that the stratigraphically higher fossils, which include the important tibia, humerus and a large, presumed male, mandible (KNM-KP 29287), might belong to a later, more derived hominid. We have recovered new fossils from Kanapoi and Allia Bay, Kenya, during field work in 1995-1997 that confirm the primitive status of Australopithecus anamensis, the earliest species of Australopithecus. Isotope dating confirms A. anamensis' intermediate age as being between those of Ardipithecus ramidus and Australopithecus afarensis. New specimens of maxilla, mandible and capitate show that this species is demonstrably more primitive than A. afarensis. A lower first deciduous molar (dm 1) is intermediate in morphology between that reported for Ardipithecus ramidus and A. afarensis. Single-crystal 40Ar-39Ar age determinations on the Kanapoi Tuff show that, except for a large mandible, all of the hominid fossils from Kanapoi are from sediments deposited between 4.17+/-0.03 and 4.07+/-0.02 million years ago.

The Oldowan Stone tool industry was named for 1.8-million-year-old (Myr) artefacts found near the bottom of Olduvai Gorge, Tanzania. Subsequent archaeological research in the Omo (Ethiopia) and Turkana (Kenya) also yielded stone tools dated to 2.3 Myr. Palaeoanthropological investigations in the Hadar region of the Awash Valley of Ethiopia, revealed Oldowan assemblages in the adjacent Gona River drainage. We conducted field work in the Gona study area of Ethiopia between 1992 and 1994 which resulted in additional archaeological discoveries as well as radioisotopic age control and a magnetic polarity stratigraphy of the Gona sequence. These occurrences are now securely dated between 2.6-2.5 Myr. The stone tools are thus the oldest known artefacts from anywhere in the world. The artefacts show surprisingly sophisticated control of stone fracture mechanics, equivalent to much younger Oldowan assemblages of Early Pleistocene age. This indicates an unexpectedly long period of technological stasis in the Oldowan.

Nine hominid dental, cranial and postcranial specimens from Kanapoi, Kenya, and 12 specimens from Allia Bay, Kenya, are described here as a new species of Australopithecus dating from between about 3.9 million and 4.2 million years ago. The mosaic of primitive and derived features shows this species to be a possible ancestor to Australopithecus afarensis and suggests that Ardipithecus ramidus is a sister species to this and all later hominids. A tibia establishes that hominids were bipedal at least half a million years before the previous earliest evidence showed.

The chronometric framework developed for Plio-Pleistocene deposits of the northern Turkana Basin is reviewed in light of recent advances in lithostratigraphy, geochemical correlation, paleomagnetic stratigraphy, and isotopic dating. The sequence is tightly controlled by 20 precise ages on volcanic materials. These ages are internally consistent but are at variance with estimates for the boundaries of the magnetic polarity time scale by about 0.07 my. This discrepancy can be only partially resolved at present. Based on the established chronometric framework and stratigraphic sequences, depositional ages can be estimated for significant marker beds. These ages can in turn be used to constrain the 449 hominid specimens thus far reported from the basin. Ages for most hominid specimens can be estimated with a precision of +/- 0.05 my. In addition, the chronometric framework will be applicable to other paleontological collections, archeological excavations, and future discoveries in the basin.