Age of the oldest known Homo sapiens from eastern Africa.

Efforts to date the oldest modern human fossils in eastern Africa, from Omo-Kibish1-3 and Herto4,5 in Ethiopia, have drawn on a variety of chronometric evidence, including 40Ar/39Ar ages of stratigraphically associated tuffs. The ages that are generally reported for these fossils are around 197 thousand years (kyr) for the Kibish Omo I3,6,7, and around 160-155 kyr for the Herto hominins5,8. However, the stratigraphic relationships and tephra correlations that underpin these estimates have been challenged6,8. Here we report geochemical analyses that link the Kamoya's Hominid Site (KHS) Tuff9, which conclusively overlies the member of the Omo-Kibish Formation that contains Omo I, with a major explosive eruption of Shala volcano in the Main Ethiopian Rift. By dating the proximal deposits of this eruption, we obtain a new minimum age for the Omo fossils of 233 ± 22 kyr. Contrary to previous arguments6,8, we also show that the KHS Tuff does not correlate with another widespread tephra layer, the Waidedo Vitric Tuff, and therefore cannot anchor a minimum age for the Herto fossils. Shifting the age of the oldest known Homo sapiens fossils in eastern Africa to before around 200 thousand years ago is consistent with independent evidence for greater antiquity of the modern human lineage10.

Age and context of mid-Pliocene hominin cranium from Woranso-Mille, Ethiopia.

A fossil hominin cranium was discovered in mid-Pliocene deltaic strata in the Godaya Valley of the northwestern Woranso-Mille study area in Ethiopia. Here we show that analyses of chemically correlated volcanic layers and the palaeomagnetic stratigraphy, combined with Bayesian modelling of dated tuffs, yield an age range of 3.804 ± 0.013 to 3.777 ± 0.014 million years old (mean ± 1σ) for the deltaic strata and the fossils that they contain. We also document deposits of a perennial lake beneath the deltaic sequence. Mammalian fossils associated with the cranium represent taxa that were widespread at the time and data from botanical remains indicate that the vegetation in the lake and delta catchment was predominantly dry shrubland with varying proportions of grassland, wetland and riparian forest. In addition, we report high rates of sediment accumulation and depositional features that are typical of a steep topographic relief and differ from younger Woranso-Mille fossil localities, reflecting the influence of active rift processes on the palaeolandscape.

Holocene bidirectional river system along the Kenya Rift and its influence on East African faunal exchange and diversity gradients.

East Africa is a global biodiversity hotspot and exhibits distinct longitudinal diversity gradients from west to east in freshwater fishes and forest mammals. The assembly of this exceptional biodiversity and the drivers behind diversity gradients remain poorly understood, with diversification often studied at local scales and less attention paid to biotic exchange between Afrotropical regions. Here, we reconstruct a river system that existed for several millennia along the now semiarid Kenya Rift Valley during the humid early Holocene and show how this river system influenced postglacial dispersal of fishes and mammals due to its dual role as a dispersal corridor and barrier. Using geomorphological, geochronological, isotopic, and fossil analyses and a synthesis of radiocarbon dates, we find that the overflow of Kenyan rift lakes between 12 and 8 ka before present formed a bidirectional river system consisting of a "Northern River" connected to the Nile Basin and a "Southern River," a closed basin. The drainage divide between these rivers represented the only viable terrestrial dispersal corridor across the rift. The degree and duration of past hydrological connectivity between adjacent river basins determined spatial diversity gradients for East African fishes. Our reconstruction explains the isolated distribution of Nilotic fish species in modern Kenyan rift lakes, Guineo-Congolian mammal species in forests east of the Kenya Rift, and recent incipient vertebrate speciation and local endemism in this region. Climate-driven rearrangements of drainage networks unrelated to tectonic activity contributed significantly to the assembly of species diversity and modern faunas in the East African biodiversity hotspot.

Plio-Pleistocene environmental variability in Africa and its implications for mammalian evolution.

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.

Comparative description and taxonomic affinity of 3.7-million-year-old hominin mandibles from Woranso-Mille (Ethiopia).

Fossil discoveries of early Australopithecus species from Woranso-Mille have played a significant role in improving our understanding of mid-Pliocene hominin evolution and diversity. Here, we describe two mandibles with dentitions, recovered from sediments immediately above a tuff radiometrically dated to 3.76 ± 0.02 Ma, and assess their taxonomic affinity. The two mandibles (MSD-VP-5/16 and MSD-VP-5/50) show morphological similarities with both Australopithecus anamensis and Australopithecus afarensis. Some of the unique features that distinguish Au. anamensis from Au. afarensis are present in the mandibles, which also share a few derived features with Au. afarensis. Their retention of more Kanapoi Au. anamensis-like traits, compared to the fewer derived features they share with Au. afarensis, and the presence of Au. anamensis at Woranso-Mille in 3.8-million-year-old deposits, lends support to their assignment to Au. anamensis. However, it is equally arguable that the few derived dentognathic features they share with Au. afarensis could be taxonomically more significant, making it difficult to conclusively assign these specimens to either species. Regardless of which species they are assigned to, the mosaic nature of the dentognathic morphology and geological age of the two mandibles lends further support to the hypothesized ancestor-descendant relationship between Au. anamensis and Au. afarensis. However, there is now limited fossil evidence indicating that these two species may have overlapped in time. Hence, the last appearance of Au. anamensis and first appearance of Au. afarensis are currently unknown. Recovery of Australopithecus fossils from 4.1 to 3.8 Ma is critical to further address the timing of these events.

Earliest known Oldowan artifacts at >2.58 Ma from Ledi-Geraru, Ethiopia, highlight early technological diversity.

The manufacture of flaked stone artifacts represents a major milestone in the technology of the human lineage. Although the earliest production of primitive stone tools, predating the genus Homo and emphasizing percussive activities, has been reported at 3.3 million years ago (Ma) from Lomekwi, Kenya, the systematic production of sharp-edged stone tools is unknown before the 2.58-2.55 Ma Oldowan assemblages from Gona, Ethiopia. The organized production of Oldowan stone artifacts is part of a suite of characteristics that is often associated with the adaptive grade shift linked to the genus Homo Recent discoveries from Ledi-Geraru (LG), Ethiopia, place the first occurrence of Homo ∼250 thousand years earlier than the Oldowan at Gona. Here, we describe a substantial assemblage of systematically flaked stone tools excavated in situ from a stratigraphically constrained context [Bokol Dora 1, (BD 1) hereafter] at LG bracketed between 2.61 and 2.58 Ma. Although perhaps more primitive in some respects, quantitative analysis suggests the BD 1 assemblage fits more closely with the variability previously described for the Oldowan than with the earlier Lomekwian or with stone tools produced by modern nonhuman primates. These differences suggest that hominin technology is distinctly different from generalized tool use that may be a shared feature of much of the primate lineage. The BD 1 assemblage, near the origin of our genus, provides a link between behavioral adaptations-in the form of flaked stone artifacts-and the biological evolution of our ancestors.

New Pliocene hominin remains from the Leado Dido'a area of Woranso-Mille, Ethiopia.

Fossiliferous deposits at Woranso-Mille span the period when Australopithecus anamensis gave rise to Australopithecus afarensis (3.8-3.6 Ma) and encompass the core of the A. afarensis range (ca. 3.5-3.2 Ma). Within the latter period, fossils described to date include the intriguing but taxonomically unattributed Burtele foot, dentognathic fossils attributed to Australopithecus deyiremeda, and one specimen securely attributed to A. afarensis (the Nefuraytu mandible). These fossils suggest that at least one additional hominin lineage lived alongside A. afarensis in the Afar Depression. Here we describe a collection of hominin fossils from a new locality in the Leado Dido'a area of Woranso-Mille (LDD-VP-1). The strata in this area are correlated to the same chron as those in the Burtele area (C2An.3n; 3.59-3.33 Ma), and similar in age to the Maka Sands and the Basal through lower Sidi Hakoma Members of the Hadar Formation. We attribute all but one of the LDD hominin specimens to A. afarensis, based on diagnostic morphology of the mandible, maxilla, canines, and premolars. The LDD specimens generally fall within the range of variation previously documented for A. afarensis but increase the frequency of some rare morphological variants. However, one isolated M3 is extremely small, and its taxonomic affinity is currently unknown. The new observations support previous work on temporal trends in A. afarensis and demonstrate that the large range of variation accepted for this species is present even within a limited spatiotemporal range. The value added with this sample lies in its contribution to controlling for spatiotemporal differences among site samples in the A. afarensis hypodigm and its contemporaneity with non-A. afarensis specimens at Woranso-Mille.

New species from Ethiopia further expands Middle Pliocene hominin diversity.

Middle Pliocene hominin species diversity has been a subject of debate over the past two decades, particularly after the naming of Australopithecus bahrelghazali and Kenyanthropus platyops in addition to the well-known species Australopithecus afarensis. Further analyses continue to support the proposal that several hominin species co-existed during this time period. Here we recognize a new hominin species (Australopithecus deyiremeda sp. nov.) from 3.3-3.5-million-year-old deposits in the Woranso-Mille study area, central Afar, Ethiopia. The new species from Woranso-Mille shows that there were at least two contemporaneous hominin species living in the Afar region of Ethiopia between 3.3 and 3.5 million years ago, and further confirms early hominin taxonomic diversity in eastern Africa during the Middle Pliocene epoch. The morphology of Au. deyiremeda also reinforces concerns related to dentognathic (that is, jaws and teeth) homoplasy in Plio-Pleistocene hominins, and shows that some dentognathic features traditionally associated with Paranthropus and Homo appeared in the fossil record earlier than previously thought.

Progressive aridification in East Africa over the last half million years and implications for human evolution.

Evidence for Quaternary climate change in East Africa has been derived from outcrops on land and lake cores and from marine dust, leaf wax, and pollen records. These data have previously been used to evaluate the impact of climate change on hominin evolution, but correlations have proved to be difficult, given poor data continuity and the great distances between marine cores and terrestrial basins where fossil evidence is located. Here, we present continental coring evidence for progressive aridification since about 575 thousand years before present (ka), based on Lake Magadi (Kenya) sediments. This long-term drying trend was interrupted by many wet-dry cycles, with the greatest variability developing during times of high eccentricity-modulated precession. Intense aridification apparent in the Magadi record took place between 525 and 400 ka, with relatively persistent arid conditions after 350 ka and through to the present. Arid conditions in the Magadi Basin coincide with the Mid-Brunhes Event and overlap with mammalian extinctions in the South Kenya Rift between 500 and 400 ka. The 525 to 400 ka arid phase developed in the South Kenya Rift between the period when the last Acheulean tools are reported (at about 500 ka) and before the appearance of Middle Stone Age artifacts (by about 320 ka). Our data suggest that increasing Middle- to Late-Pleistocene aridification and environmental variability may have been drivers in the physical and cultural evolution of Homo sapiens in East Africa.

Evidence for rapid faunal change in the early Miocene of East Africa based on revised biostratigraphic and radiometric dating of Bukwa, Uganda.

Field expeditions to Bukwa in the late 1960s and early 1970s established that the site had a small but diverse early Miocene fauna, including the catarrhine primate Limnopithecus legetet. Initial potassium-argon radiometric dating indicated that Bukwa was 22 Ma, making it the oldest of the East African early Miocene fossil localities known at the time. In contrast, the fauna collected from Bukwa was similar to other fossil localities in the region that were several million years younger. This discrepancy was never resolved, and due to the paucity of primate remains at the site, little subsequent research took place. We have collected new fossils at Bukwa, reanalyzed the existing fossil collections, and provided new radiometric dating. 40Ar/39Ar incremental heating ages on lavas bracketing the site indicate that the Bukwa fossils were deposited ∼19 Ma, roughly 3 Ma younger than the original radiometric age. Our radiometric dating results are corroborated by a thorough reanalysis of the faunal assemblage. Bukwa shares taxa with both stratigraphically older localities (Tinderet, Napak) and with stratigraphically younger localities (Kisingiri, Turkana Basin) perfectly corresponding to our revised radiometric age. This revised age for Bukwa is important because it indicates that significant faunal turnover may have occurred in East Africa between 20 and 19 Ma. Bukwa samples immigrant taxa such as large suids, large ruminants, and ochotonids that are absent from stratigraphically older but well-sampled localities in the region, such as Tinderet (∼20 Ma) and Napak (20 Ma). Further age refinements for Bukwa and the entire East African early Miocene sequence will help to constrain the timing of this faunal turnover event, of particular importance in paleoanthropology since this temporal sequence also provides us with what is currently our best window into the early evolution of cercopithecoid and hominoid primates.

A new hominin foot from Ethiopia shows multiple Pliocene bipedal adaptations.

A newly discovered partial hominin foot skeleton from eastern Africa indicates the presence of more than one hominin locomotor adaptation at the beginning of the Late Pliocene epoch. Here we show that new pedal elements, dated to about 3.4 million years ago, belong to a species that does not match the contemporaneous Australopithecus afarensis in its morphology and inferred locomotor adaptations, but instead are more similar to the earlier Ardipithecus ramidus in possessing an opposable great toe. This not only indicates the presence of more than one hominin species at the beginning of the Late Pliocene of eastern Africa, but also indicates the persistence of a species with Ar. ramidus-like locomotor adaptation into the Late Pliocene.

Continuous 1.3-million-year record of East African hydroclimate, and implications for patterns of evolution and biodiversity.

The transport of moisture in the tropics is a critical process for the global energy budget and on geologic timescales, has markedly influenced continental landscapes, migratory pathways, and biological evolution. Here we present a continuous, first-of-its-kind 1.3-My record of continental hydroclimate and lake-level variability derived from drill core data from Lake Malawi, East Africa (9-15° S). Over the Quaternary, we observe dramatic shifts in effective moisture, resulting in large-scale changes in one of the world's largest lakes and most diverse freshwater ecosystems. Results show evidence for 24 lake level drops of more than 200 m during the Late Quaternary, including 15 lowstands when water levels were more than 400 m lower than modern. A dramatic shift is observed at the Mid-Pleistocene Transition (MPT), consistent with far-field climate forcing, which separates vastly different hydroclimate regimes before and after ∼800,000 years ago. Before 800 ka, lake levels were lower, indicating a climate drier than today, and water levels changed frequently. Following the MPT high-amplitude lake level variations dominate the record. From 800 to 100 ka, a deep, often overfilled lake occupied the basin, indicating a wetter climate, but these highstands were interrupted by prolonged intervals of extreme drought. Periods of high lake level are observed during times of high eccentricity. The extreme hydroclimate variability exerted a profound influence on the Lake Malawi endemic cichlid fish species flock; the geographically extensive habitat reconfiguration provided novel ecological opportunities, enabling new populations to differentiate rapidly to distinct species.

Tephrostratigraphy of the Waki-Mille area of the Woranso-Mille paleoanthropological research project, Afar, Ethiopia.

Tephra geochemistry and (40)Ar/(39)Ar geochronology are reported for the Waki-Mille area in the northwestern part of the Woranso-Mille paleoanthropological project area in the west central Afar region of Ethiopia. Previous studies documented dentognathic fossils that are morphologically intermediate between Australopithecus anamensis and Australopithecus afarensis and some that are attributed to Australopithecus afarensis. Additional dentognathic remains from the study area were assigned to the newly identified species Australopithecus deyiremeda. These fossil hominin taxa were recovered from volcanic and sedimentary strata containing tuffs ranging in age from more than 3.77 million years ago (Ma) to less than 3.469 Ma. One of the tuffs was correlated based on geochemistry, feldspar mineralogy, and age to the Lokochot Tuff of the Omo-Turkana Basin of southern Ethiopia and Kenya. Variations in major and minor element abundances in volcanic glass demarcate ten geochemically distinct tuffs and tuff sequences, including three that are geochemically similar to widespread regional tuffs, specifically the Lomogol, Lokochot, and ß- Tulu Bor/Sidi Hakoma tuffs. A new (40)Ar/(39)Ar age for the Waki Tuff, which is geochemically similar to the Lomogol Tuff, is 3.664 ± 0.016 Ma. Other tuffs in the Waki-Mille area are geochemically dissimilar to regional tuffs documented to date. Identification of tuffs based on character, stratigraphic position, and geochemistry refines local stratigraphic correlations and delineates the geographic distributions of precisely dated fossiliferous levels within the Waki-Mille area.

Dentognathic remains of Australopithecus afarensis from Nefuraytu (Woranso-Mille, Ethiopia): Comparative description, geology, and paleoecological context.

Australopithecus afarensis is the best-known and most dimorphic species in the early hominin fossil record. Here, we present a comparative description of new fossil specimens of Au. afarensis from Nefuraytu, a 3.330-3.207 million-years-old fossil collection area in the Woranso-Mille study area, central Afar, Ethiopia. These specimens include NFR-VP-1/29, one of the most complete mandibles assigned to the species thus far and among the largest mandibles attributed to Au. afarensis, likely representing a male individual. NFR-VP-1/29 retains almost all of the distinctive archaic features documented for Au. afarensis. These features include a posteriorly sloping symphysis, a low and rounded basally set inferior transverse torus, anterosuperiorly opening mental foramen, a lateral corpus hollow bound anteriorly by the C/P3 jugae and posteriorly by the lateral prominence, and the ascending ramus arising high on the corpus. Dental morphology and metrics of the Nefuraytu specimens also falls within the range of Au. afarensis. The presence of this species at Woranso-Mille between 3.330 and 3.207 million years ago confirms the existence of this species in the area in close spatial and temporal proximity to other middle Pliocene hominin taxa such as the one represented by the Burtele foot (BRT-VP-2/73) and the recently named species Australopithecus deyiremeda. This has important implications for our understanding of middle Pliocene hominin diversity.

An early Australopithecus afarensis postcranium from Woranso-Mille, Ethiopia.

Only one partial skeleton that includes both forelimb and hindlimb elements has been reported for Australopithecus afarensis. The diminutive size of this specimen (A.L. 288-1 ["Lucy"]) has hampered our understanding of the paleobiology of this species absent the potential impact of allometry. Here we describe a large-bodied (i.e., well within the range of living Homo) specimen that, at 3.58 Ma, also substantially antedates A.L. 288-1. It provides fundamental evidence of limb proportions, thoracic form, and locomotor heritage in Australopithecus afarensis. Together, these characteristics further establish that bipedality in Australopithecus was highly evolved and that thoracic form differed substantially from that of either extant African ape.

Oldest evidence of abundant C4 grasses and habitat heterogeneity in eastern Africa.

The assembly of Africa's iconic C4 grassland ecosystems is central to evolutionary interpretations of many mammal lineages, including hominins. C4 grasses are thought to have become ecologically dominant in Africa only after 10 million years ago (Ma). However, paleobotanical records older than 10 Ma are sparse, limiting assessment of the timing and nature of C4 biomass expansion. This study uses a multiproxy design to document vegetation structure from nine Early Miocene mammal site complexes across eastern Africa. Results demonstrate that between ~21 and 16 Ma, C4 grasses were locally abundant, contributing to heterogeneous habitats ranging from forests to wooded grasslands. These data push back the oldest evidence of C4 grass-dominated habitats in Africa-and globally-by more than 10 million years, calling for revised paleoecological interpretations of mammalian evolution.

The evolution of hominoid locomotor versatility: Evidence from Moroto, a 21 Ma site in Uganda.

Living hominoids are distinguished by upright torsos and versatile locomotion. It is hypothesized that these features evolved for feeding on fruit from terminal branches in forests. To investigate the evolutionary context of hominoid adaptive origins, we analyzed multiple paleoenvironmental proxies in conjunction with hominoid fossils from the Moroto II site in Uganda. The data indicate seasonally dry woodlands with the earliest evidence of abundant C4 grasses in Africa based on a confirmed age of 21 million years ago (Ma). We demonstrate that the leaf-eating hominoid Morotopithecus consumed water-stressed vegetation, and postcrania from the site indicate ape-like locomotor adaptations. These findings suggest that the origin of hominoid locomotor versatility is associated with foraging on leaves in heterogeneous, open woodlands rather than forests.

(40)Ar/(39)Ar dating of Bed I, Olduvai Gorge, Tanzania, and the chronology of early Pleistocene climate change.

(40)Ar/(39)Ar dating of tuffs and lavas of the late Pleistocene volcanic and sedimentary sequence of Olduvai Gorge, north-central Tanzania, provides the basis for a revision of Bed I chronostratigraphy. Bed I extends from immediately above the Naabi Ignimbrite at 2.038 ± 0.005 Ma to Tuff IF at 1.803 ± 0.002 Ma. Tuff IB, a prominent widespread marker tuff in the basin and a key to understanding hominin evolutionary chronologies and paleoclimate histories, has an age of 1.848 ± 0.003 Ma. The largest lake expansion event in the closed Olduvai lake basin during Bed I times encompassed the episode of eruption and emplacement of this tuff. This lake event is nearly coincident with the maximum precessional insolation peak of the entire Bed I/Lower Bed II interval, calculated from an astronomical model of the boreal summer orbital insolation time-series. The succeeding precessional peak also apparently coincides with the next youngest expansion of paleo-Lake Olduvai. The extreme wet/dry climate shifts seen in the upper part of Bed I occur during an Earth-orbital eccentricity maximum, similar to episodic lake expansions documented elsewhere in the East African Rift during the Neogene.