Naomi E. Levin

High-temperature environments of human evolution in East Africa based on bond ordering in paleosol carbonates

Many important hominid-bearing fossil localities in East Africa are in regions that are extremely hot and dry. Although humans are well adapted to such conditions, it has been inferred that East African environments were cooler or more wooded during the Pliocene and Pleistocene when this region was a central stage of human evolution. Here we show that the Turkana Basin, Kenya—today one of the hottest places on Earth—has been continually hot during the past 4 million years. The distribution of 13C-18O bonds in paleosol carbonates indicates that soil temperatures during periods of carbonate formation were typically above 30 °C and often in excess of 35 °C. Similar soil temperatures are observed today in the Turkana Basin and reflect high air temperatures combined with solar heating of the soil surface. These results are specific to periods of soil carbonate formation, and we suggest that such periods composed a large fraction of integrated time in the Turkana Basin. If correct, this interpretation has implications for human thermophysiology and implies a long-standing human association with marginal environments.

Woody cover and hominin environments in the past 6 million years

The role of African savannahs in the evolution of early hominins has been debated for nearly a century. Resolution of this issue has been hindered by difficulty in quantifying the fraction of woody cover in the fossil record. Here we show that the fraction of woody cover in tropical ecosystems can be quantified using stable carbon isotopes in soils. Furthermore, we use fossil soils from hominin sites in the Awash and Omo-Turkana basins in eastern Africa to reconstruct the fraction of woody cover since the Late Miocene epoch (about 7 million years ago). 13C/12C ratio data from 1,300 palaeosols at or adjacent to hominin sites dating to at least 6 million years ago show that woody cover was predominantly less than ∼40% at most sites. These data point to the prevalence of open environments at the majority of hominin fossil sites in eastern Africa over the past 6 million years.

A Female Homo erectus Pelvis from Gona, Ethiopia

Analyses of the KNM-WT 15000 Homo erectus juvenile male partial skeleton from Kenya concluded that this species had a tall thin body shape due to specialized locomotor and climatic adaptations. Moreover, it was concluded that H. erectus pelves were obstetrically restricted to birthing a small-brained altricial neonate. Here we describe a nearly complete early Pleistocene adult female H. erectus pelvis from the Busidima Formation of Gona, Afar, Ethiopia. This obstetrically capacious pelvis demonstrates that pelvic shape in H. erectus was evolving in response to increasing fetal brain size. This pelvis indicates that neither adaptations to tropical environments nor endurance running were primary selective factors in determining pelvis morphology in H. erectus during the early Pleistocene.

Isotopic evidence for Plio-Pleistocene environmental change at Gona, Ethiopia

Abstract A 4.5 Ma record of fluvial and lacustrine deposits is well exposed at Gona, in the Afar Depression of Ethiopia. We use isotopic values of pedogenic carbonate and fossil teeth to reconstruct Plio–Pleistocene environmental change at Gona. An increase in δ13C values of pedogenic carbonates since 4.5 Ma points to a shift from woodlands to grassy woodlands in the early Pliocene, −10.4 to −3.9‰ (VPDB), to more open but still mixed environments in the late Pleistocene, −3.0 to −1.4‰ (VPDB). This pattern is also seen in isotopic records elsewhere in East Africa. However, at 1.5 Ma the higher proportion of C4 grasses at Gona is largely a result of a local facies shift to more water-limited environments. The wide range of δ13C values of pedogenic carbonate within single stratigraphic levels indicates a mosaic of vegetation for all time intervals at Gona that depends on depositional environment. Elements of this mosaic are reflected in δ13C values of both modern plants and soil organic matter and Plio–Pleistocene soil carbonate, indicating higher amounts of C4 grasses with greater distance from a river channel in both the modern and ancient Awash River systems. δ18O values of pedogenic carbonates increase up-section from −11.9‰ in the early Pliocene to −6.4‰ (VPDB) in the late Pleistocene. The wide range of δ18O values in paleovertisol carbonates from all stratigraphic levels probably reflects short-term climate changes and periods of strong evaporation throughout the record. Based on the comparison between δ18O values of Plio–Pleistocene pedogenic carbonates and modern waters, we estimate that there has been a 6.5‰ increase in mean annual δ18O values of meteoric water since 4.5 Ma. δ18O values of pedogenic carbonate from other East African records indicate a similar shift. Increasing aridity and fluctuations in the timing and source of rainfall are likely responsible for the changes in δ18O values of East African pedogenic carbonates through the Plio–Pleistocene.

Early hominin diet included diverse terrestrial and aquatic animals 1.95 Ma in East Turkana, Kenya

The manufacture of stone tools and their use to access animal tissues by Pliocene hominins marks the origin of a key adaptation in human evolutionary history. Here we report an in situ archaeological assemblage from the Koobi Fora Formation in northern Kenya that provides a unique combination of faunal remains, some with direct evidence of butchery, and Oldowan artifacts, which are well dated to 1.95 Ma. This site provides the oldest in situ evidence that hominins, predating Homo erectus, enjoyed access to carcasses of terrestrial and aquatic animals that they butchered in a well-watered habitat. It also provides the earliest definitive evidence of the incorporation into the hominin diet of various aquatic animals including turtles, crocodiles, and fish, which are rich sources of specific nutrients needed in human brain growth. The evidence here shows that these critical brain-growth compounds were part of the diets of hominins before the appearance of Homo ergaster/erectus and could have played an important role in the evolution of larger brains in the early history of our lineage.

Early Pliocene hominids from Gona, Ethiopia

Comparative biomolecular studies suggest that the last common ancestor of humans and chimpanzees, our closest living relatives, lived during the Late Miocene–Early Pliocene. Fossil evidence of Late Miocene–Early Pliocene hominid evolution is rare and limited to a few sites in Ethiopia, Kenya and Chad. Here we report new Early Pliocene hominid discoveries and their palaeoenvironmental context from the fossiliferous deposits of As Duma, Gona Western Margin (GWM), Afar, Ethiopia. The hominid dental anatomy (occlusal enamel thickness, absolute and relative size of the first and second lower molar crowns, and premolar crown and radicular anatomy) indicates attribution to Ardipithecus ramidus. The combined radioisotopic and palaeomagnetic data suggest an age of between 4.51 and 4.32 million years for the hominid finds at As Duma. Diverse sources of data (sedimentology, faunal composition, ecomorphological variables and stable carbon isotopic evidence from the palaeosols and fossil tooth enamel) indicate that the Early Pliocene As Duma sediments sample a moderate rainfall woodland and woodland/grassland.

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.

Northeast African vegetation change over 12 m.y.

Intense debate surrounds the evolution of grasses using the C 4 (Hatch-Slack) photosynthesis pathway and the emergence of African grasslands, often assumed to be one and the same. Here, we bring new insights with the combination of plant leaf wax carbon isotopic composition (δ 13 C wax ) and pollen data from marine sediments of the Gulf of Aden (northeast Africa), which show that C 4 biomass increases were not necessarily associated with regional grassland expansion. We find broadly opposing trends toward more enriched δ 13 C wax values and decreased grass pollen proportions between 12 and 1.4 Ma. This apparently contradictory evidence can be reconciled if a greater proportion of the Late Miocene northeast African landscape were covered by C 3 grasses than previously thought, such that C 4 grasses and shrubs replaced a C 3 ecosystem including trees and productive grasslands. In addition, δ 13 C wax and pollen both indicate that true rainforests were unlikely to have been extensive in northeast Africa at any time in the last 12 m.y., although seasonally dry forests were a significant component of the regional landscape since the Late Miocene. Here, we extend regionally integrative marine archives of terrestrial vegetation back to 12 Ma, and we evaluate them in the context of an updated compilation of pedogenic carbonate δ 13 C values from East African Rift strata. We identify two distinct phases of increasing C 4 biomass between 11 and 9 Ma (with a reversal by 4.3 Ma) and then a re-expansion between 4.3 and 1.4 Ma; surprisingly, neither was associated with grassland expansion.

Paleoenvironments of the earliest stone toolmakers, Gona, Ethiopia

Fluvio-lacustrine sediments of the Hadar and Busidima Formations along the northern Awash River (Ethiopia) archive almost three million years (3.4 to <0.6 Ma) of human evolution, including the earliest documented record of stone toolmaking at 2.5–2.6 Ma. This paper brings together sedimentologic and isotopic evidence for the paleoenvironmental context of early hominids from both formations, but with particular emphasis on the setting for the early toolmakers. The pre–2.92 Ma record (Hadar Formation) is characterized by low-gradient fl uvial, paludal, and lacustrine deposition in an undissected topography most analogous to reaches of the modern middle Awash River near Gewane. The Gona area experienced repeated deep dissection and aggradation by the Awash River, starting between 2.92 and ca. 2.7 Ma and continuing through the top of the record at <0.6 Ma (Busidima Formation). Each aggradational succession is 10–20 m in thickness and fi nes upward from wellrounded conglomerates at the base to capping paleosols at the top. During this period the ancestral Awash represented by these fi ning upward sequences was dominantly meandering and fl owed northeast, as it does today. Smaller channels tributary to the axial Awash system are also extensively exposed in the Busidima Formation. Compared to the axial-system conglomerates, the tributary channels transported fi ner, less mature volcanic clasts mixed with abundant carbonate nodules reworked from adjacent badlands. Stone artifacts (Oldowan; 2.6–2.0 Ma) at the oldest archaeological sites are only associated with the axial Awash system, in the bedded silts or capping paleosols of the fi ning upward sequences. The implements were made from rounded cobbles from the channels, but manufacture and use of the tools was always away from the channel bars, on the nearby sandy banks and silt-dominated fl oodplains. Archaeological sites higher in the record (Acheulian; <1.7 Ma) occur in similar axial river contexts, as well as along tributary channels further removed from artifact raw material sources. Mature paleosols in the Hadar and Busidima Formations are mostly pale to darkbrown Vertisols typifi ed by abundant clay slickensides, pseudo-anticlinal and vertical fracturing, and carbonate nodules. Such calcic Vertisols are common in the region today, demonstrating that the paleoclimate over the past 3.4 m.y. has been semi-arid and strongly seasonal. Carbon isotopic results from pedogenic carbonates in the Vertisols allow reconstruction of the proportion of C 3 plants (trees and shrubs) to C 4 plants (grasses) through time. The δ 13

Dietary changes of large herbivores in the Turkana Basin, Kenya from 4 to 1 Ma

Significance Stable carbon isotopes give diet information for both modern and fossil mammals and can be used to classify diets as C4 grazers, C3–C4 mixed, or C3 browsers. We show that diets of some major African herbivore lineages have significantly changed over the past 4 million years by comparing fossils from the Turkana Basin in Kenya with modern mammals from East and Central Africa. Some fossil assemblages have no modern analogues in East and Central Africa, suggesting different ecological functions for some mammals in the past as compared with their modern counterparts. The development of modern tropical grassland ecosystems are products of the coevolution of both grasses and herbivores. A large stable isotope dataset from East and Central Africa from ca. 30 regional collection sites that range from forest to grassland shows that most extant East and Central African large herbivore taxa have diets dominated by C4 grazing or C3 browsing. Comparison with the fossil record shows that faunal assemblages from ca. 4.1–2.35 Ma in the Turkana Basin had a greater diversity of C3–C4 mixed feeding taxa than is presently found in modern East and Central African environments. In contrast, the period from 2.35 to 1.0 Ma had more C4-grazing taxa, especially nonruminant C4-grazing taxa, than are found in modern environments in East and Central Africa. Many nonbovid C4 grazers became extinct in Africa, notably the suid Notochoerus, the hipparion equid Eurygnathohippus, the giraffid Sivatherium, and the elephantid Elephas. Other important nonruminant C4-grazing taxa switched to browsing, including suids in the lineage Kolpochoerus-Hylochoerus and the elephant Loxodonta. Many modern herbivore taxa in Africa have diets that differ significantly from their fossil relatives. Elephants and tragelaphin bovids are two groups often used for paleoecological insight, yet their fossil diets were very different from their modern closest relatives; therefore, their taxonomic presence in a fossil assemblage does not indicate they had a similar ecological function in the past as they do at present. Overall, we find ecological assemblages of C3-browsing, C3–C4-mixed feeding, and C4-grazing taxa in the Turkana Basin fossil record that are different from any modern ecosystem in East or Central Africa.