The age of the hominin fossils from Jebel Irhoud, Morocco, and the origins of the Middle Stone Age.

The timing and location of the emergence of our species and of associated behavioural changes are crucial for our understanding of human evolution. The earliest fossil attributed to a modern form of Homo sapiens comes from eastern Africa and is approximately 195 thousand years old, therefore the emergence of modern human biology is commonly placed at around 200 thousand years ago. The earliest Middle Stone Age assemblages come from eastern and southern Africa but date much earlier. Here we report the ages, determined by thermoluminescence dating, of fire-heated flint artefacts obtained from new excavations at the Middle Stone Age site of Jebel Irhoud, Morocco, which are directly associated with newly discovered remains of H. sapiens. A weighted average age places these Middle Stone Age artefacts and fossils at 315 ± 34 thousand years ago. Support is obtained through the recalculated uranium series with electron spin resonance date of 286 ± 32 thousand years ago for a tooth from the Irhoud 3 hominin mandible. These ages are also consistent with the faunal and microfaunal assemblages and almost double the previous age estimates for the lower part of the deposits. The north African site of Jebel Irhoud contains one of the earliest directly dated Middle Stone Age assemblages, and its associated human remains are the oldest reported for H. sapiens. The emergence of our species and of the Middle Stone Age appear to be close in time, and these data suggest a larger scale, potentially pan-African, origin for both.

Perissodactyla (Rhinocerotidae and Equidae) from Kanapoi.

The Kanapoi collection of Rhinocerotidae, first studied by Hooijer and Patterson (1972), now consists of 25 specimens and substantial reinterpretation of their affinities is made here. Kanapoi post-dates the extinction of Brachypotherium and the whole collection belongs to the Dicerotini. It is important because it includes the type-specimen of Diceros praecox, a species that remains poorly known, but looks slightly larger and more primitive than the modern 'black' rhino, Diceros bicornis. A second species is probably ancestral to the modern 'white' rhino, Ceratotherium simum; it looks identical to the Pleistocene North African Ceratotherium mauritanicum, of which Ceratotherium efficax is probably a synonym. The evolution of the Dicerotini in Africa can be regarded as an increasing divergence in diet and related morphofunctional adaptations in the two lineages. The co-occurrence at Kanapoi of both Diceros and Ceratotherium, with distinct dietary preferences, suggests some habitat heterogeneity, although the low sample size prevents robust paleoecological conclusions. The Equidae are also rare and consist mostly of isolated teeth. I take the most parsimonious option of tentatively including all of them in a single species, whose identification is left open. Dental features of eastern African Pliocene to Pleistocene hipparions may reflect increasing adaptation to grazing.

Kantis: A new Australopithecus site on the shoulders of the Rift Valley near Nairobi, Kenya.

Most Plio-Pleistocene sites in the Gregory Rift Valley that have yielded abundant fossil hominins lie on the Rift Valley floor. Here we report a new Pliocene site, Kantis, on the shoulder of the Gregory Rift Valley, which extends the geographical range of Australopithecus afarensis to the highlands of Kenya. This species, known from sites in Ethiopia, Tanzania, and possibly Kenya, is believed to be adapted to a wide spectrum of habitats, from open grassland to woodland. The Kantis fauna is generally similar to that reported from other contemporaneous A. afarensis sites on the Rift Valley floor. However, its faunal composition and stable carbon isotopic data from dental enamel suggest a stronger C4 environment than that present at those sites. Although the Gregory Rift Valley has been the focus of paleontologists' attention for many years, surveys of the Rift shoulder may provide new perspective on African Pliocene mammal and hominin evolution.

The Omo Mursi Formation: a window into the East African Pliocene.

Dating to more than four million years ago (Ma), the Mursi Formation is among the oldest of the Plio-Pleistocene Omo Group deposits in the lower Omo Valley of southwestern Ethiopia. The sedimentary sequence is exposed along a strip ∼35 km by 4 km, but it has received relatively little attention due to the difficult access to this area. Although expeditions to the lower Omo Valley between 1968 and 1973 focused primarily on the Usno and Shungura Formations, survey of the Mursi Formation produced a faunal collection of about 250 specimens deriving exclusively from the Yellow Sands area at the southern extent of the exposures. In 2009, we reinitiated an investigation of the formation by focusing on the most northern exposures, and a new fossil site, Cholo, was identified. Cholo is depositionally similar to the lowermost exposures at the Yellow Sands, although no stratigraphic correlation between the two localities has yet been made. The fossiliferous sediments at Cholo are capped by a prominent vitric tuff that is compositionally distinct from any other known tephra preserved in East African rift basins, including the only known vitric tuff at the Yellow Sands. The faunal assemblage of the Yellow Sands area presents interesting characteristics: the fossils generally show little weathering and include a large proportion of suids (44% of the mammalian fauna) and a small proportion of bovids (14%) compared with other Pliocene African sites. The sample is also unusual in the high frequency of deinotheres (7%). Taxon-specific stable carbon isotopic composition of the Mursi mammals tends to show generally higher proportions of C3 diets compared with other Pliocene sites in East Africa and Chad. This and the particular faunal proportions suggest that the environments represented by the Mursi Formation were more closed than those of other Pliocene sites.

The Pleistocene high-elevation environments between 2.02 and 0.6 Ma at Melka Kunture (Upper Awash Valley, Ethiopia) based upon stable isotope analysis.

Pleistocene environments are among the most studied issues in paleoecology and human evolution research in eastern Africa. Many data have been recorded from archaeological sites located at low and medium elevations (≤ 1500 m), whereas few contexts are known at 2000 m and above. Here, we present a substantial isotopic study from Melka Kunture, a complex of prehistoric sites located at 2000-2200 m above sea level in the central Ethiopian highlands. We analyzed the stable carbon and oxygen isotopic composition of 308 faunal tooth enamel samples from sites dated between 2.02 and 0.6 Ma to investigate the animal diets and habitats. The carbon isotopic results indicate that the analyzed taxa had C4-dominated and mixed C3-C4 diets with no significant diachronic changes in feeding behavior with time. This is consistent with faunal and phytolith analyses, which suggested environments characterized by open grasslands (with both C3 and C4 grasses), patches of bushes and thickets, and aquatic vegetation. However, palynological data previously documented mountain forests, woodlands, and high-elevation grasslands. Additionally, the carbon isotopic comparison with other eastern African localities shows that differences in elevation did not influence animal feeding strategies and habitat partitioning, even though plant species vary according to altitudinal gradients. In contrast, the oxygen isotopic comparison suggests significant differences consistent with the altitude effect. Our approach allows us to detect diverse aspects of animal behavior, habitat, and vegetation that should be considered when reconstructing past environments.

Dietary and paleoenvironmental reconstruction using stable isotopes of herbivore tooth enamel from middle Pliocene Dikika, Ethiopia: implication for Australopithecus afarensis habitat and food resources.

Carbon and oxygen isotopes of mammalian tooth enamel were used to reconstruct paleoenvironments of Australopithecus afarensis from the middle Pliocene locality of Dikika, Ethiopia. Isotopic analyses were conducted on 210 mammalian herbivore teeth from 15 different taxa collected from the Basal Member (~3.8-3.42 Ma) and Sidi Hakoma Member (3.42-3.24 Ma) of the Hadar Formation. The isotopic analyses aim specifically at reconstructing shifts in the relative abundance of C(4) grasses in mammalian diets, and more generally at paleoclimate factors such as aridity and seasonality, as well as habitat structure. Carbon isotopic data suggest a wide range of foraging strategies, characterized by mixed C(3)/C(4) to C(4)-dominated diets in wooded grasslands to open woodlands. Weighted average C(4) dietary proportions range between 60% and 86% in the Basal Member and 49% and 74% in the Sidi Hakoma Member. Paleoclimatic conditions based on the reconstructed mean annual water deficit from the δ(18)O(enamel) values indicate a wetter climate as compared to either the early Pliocene or the Pleistocene nearby. The middle Pliocene habitat structure at Dikika could be as diverse as open grassland and wooded grassland, and woodland to forest in the Sidi Hakoma Member while wooded grassland, woodland to grassland are evident in the Basal Member. All habitats except closed woodland and forest are persistent through both members; however, the relative proportion of individual habitats changed through time. These changes could have put the fauna in competition for preferred habitats and food resources, which could have forced migration, adaptation to other resources and/or extinction. Thus, the existence of A. afarensis throughout the middle Pliocene indicates either this species might have adapted to a wide range of habitats, or its preferred habitat was not affected by the observed environmental changes.

Early Homo erectus lived at high altitudes and produced both Oldowan and Acheulean tools.

In Africa, the scarcity of hominin remains found in direct association with stone tools has hindered attempts to link Homo habilis and Homo erectus with particular lithic industries. The infant mandible discovered in level E at Garba IV (Melka Kunture) on the highlands of Ethiopia is critical to this issue due to its direct association with an Oldowan lithic industry. Here, we use synchrotron imaging to examine the internal morphology of the unerupted permanent dentition and confirm its identification as Homo erectus. Additionally, we utilize new palaeomagnetic ages to show that (i) the mandible in level E is ca. 2 million-years-old, and represents one of the earliest Homo erectus fossils, and (ii) that overlying level D, ca. 1.95 million-years-old, contains the earliest known Acheulean assemblage.

From leaves to seeds? The dietary shift in late Miocene colobine monkeys of southeastern Europe.

Extant colobine monkeys are specialized leaf eaters. But during the late Miocene, western Eurasia was home to colobines that were less efficient at chewing leaves than they were at breaking seed shells. To understand the link between folivory and granivory in this lineage, the dietary niche of Mesopithecus delsoni and Mesopithecus pentelicus was investigated in southeastern Europe, where a major environmental change occurred during the late Miocene. We combined dental topographic estimates of chewing efficiency with dental microwear texture analysis of enamel wear facets. Mesopithecus delsoni was more efficient at chewing leaves than M. pentelicus, the dental topography of which matches an opportunistic seed eater. Concurrently, microwear complexity increases in M. pentelicus, especially in the northernmost localities corresponding to present-day Bulgaria. This is interpreted as a dietary shift toward hard foods such as seeds or tubers, which is consistent with the savanna and open mixed forest biomes that covered Bulgaria during the Tortonian. The fact that M. delsoni was better adapted to folivory and consumed a lower amount of hard foods than M. pentelicus suggests that colobines either adapted to folivory before their dispersal to Europe or evolved adaptations to leaf consumption in multiple occurrences.

Earliest African evidence of carcass processing and consumption in cave at 700 ka, Casablanca, Morocco.

To date, in Africa, evidence for animal processing and consumption in caves routinely used as living spaces is only documented in the late Middle Pleistocene of the North and South of the continent and postdates the Middle Pleistocene in East Africa. Here we report the earliest evidence in a North-African cave (Grotte des Rhinocéros at Casablanca, Morocco) of cut, percussion and human gnawing marks on faunal remains directly associated with lithic knapping activities in the same space and in a well-documented stratified context. Ages for this Acheulean site are provided by the dating of herbivorous teeth to 690-720 ka and 520-550 ka (lower and upper sets) by combined Electron Spin Resonance (ESR) and U-series techniques. Traces of butchery on gazelle, alcelaphin, and zebra bones demonstrate that hominins had primary access to herbivore carcasses. Hominins brought and consumed meat in the cave, as documented by herbivore bones bearing human tooth marks concentrated in a circumscribed area of the excavation. In Africa, this site provides the earliest evidence for in situ carcass processing and meat-eating in cave, directly associated with lithic production and demonstrates the recurrent use by early Middle Pleistocene hominins of a North African cave site 400 000 years before that by Homo sapiens at Jebel Irhoud (Morocco).

Folivory or fruit/seed predation for Mesopithecus, an earliest colobine from the late Miocene of Eurasia?

Here we compare dental microwear textures from specimens of the fossil genus Mesopithecus (Cercopithecidae, Colobinae) from the late Miocene of Eastern Europe with dental microwear textures from four extant primate species with known dietary differences. Results indicate that the dental microwear textures of Mesopithecus differ from those of extant leaf eaters Alouatta palliata and Trachypithecus cristatus and instead resemble more closely those of the occasional hard-object feeders Cebus apella and Lophocebus albigena. Microwear texture data presented here in combination with results from previous analyses suggest that Mesopithecus was a widespread, opportunistic feeder that often consumed hard seeds. These data are consistent with the hypothesis that early colobines may have preferred hard seeds to leaves.

Messinian age and savannah environment of the possible hominin Graecopithecus from Europe.

Dating fossil hominids and reconstructing their environments is critically important for understanding human evolution. Here we date the potentially oldest hominin, Graecopithecus freybergi from Europe and constrain the environmental conditions under which it thrived. For the Graecopithecus-bearing Pikermi Formation of Attica/Greece, a saline aeolian dust deposit of North African (Sahara) provenance, we obtain an age of 7.37-7.11 Ma, which is coeval with a dramatic cooling in the Mediterranean region at the Tortonian-Messinian transition. Palaeobotanic proxies demonstrate C4-grass dominated wooded grassland-to-woodland habitats of a savannah biome for the Pikermi Formation. Faunal turnover at the Tortonian-Messinian transition led to the spread of new mammalian taxa along with Graecopithecus into Europe. The type mandible of G. freybergi from Pyrgos (7.175 Ma) and the single tooth (7.24 Ma) from Azmaka (Bulgaria) represent the first hominids of Messinian age from continental Europe. Our results suggest that major splits in the hominid family occurred outside Africa.

Pleistocene Hominins as a Resource for Carnivores: A c. 500,000-Year-Old Human Femur Bearing Tooth-Marks in North Africa (Thomas Quarry I, Morocco).

In many Middle Pleistocene sites, the co-occurrence of hominins with carnivores, who both contributed to faunal accumulations, suggests competition for resources as well as for living spaces. Despite this, there is very little evidence of direct interaction between them to-date. Recently, a human femoral diaphysis has been recognized in South-West of Casablanca (Morocco), in the locality called Thomas Quarry I. This site is famous for its Middle Pleistocene fossil hominins considered representatives of Homo rhodesiensis. The bone was discovered in Unit 4 of the Grotte à Hominidés (GH), dated to c. 500 ky and was associated with Acheulean artefacts and a rich mammalian fauna. Anatomically, it fits well within the group of known early Middle Pleistocene Homo, but its chief point of interest is that the diaphyseal ends display numerous tooth marks showing that it had been consumed shortly after death by a large carnivore, probably a hyena. This bone represents the first evidence of consumption of human remains by carnivores in the cave. Whether predated or scavenged, this chewed femur indicates that humans were a resource for carnivores, underlining their close relationships during the Middle Pleistocene in Atlantic Morocco.

Structure and growth pattern of pseudoteeth in Pelagornis mauretanicus (Aves, Odontopterygiformes, Pelagornithidae).

The extinct Odontopterygiformes are the sole birds known to possess strong and sharp bony pseudoteeth, the shape and location of which are closely mimetic of real teeth. The structure of the pseudoteeth is investigated here in a late Pliocene/early Pleistocene species, Pelagornis mauretanicus, using X-ray microtomography and thin sections. The results are interpreted with regard to the pseudotooth mode of growth, and have implications concerning aspects of Pelagornis ecology. The larger pseudoteeth are hollow and approximately cone-shaped, and the smaller ones are rostro-caudally constricted. The walls of pseudoteeth are composed of bone tissue of the fibro-lamellar type, which is intensively remodeled by Haversian substitution. The jaw bones display the same structure as the pseudoteeth, but their vascular canals are oriented parallel to the long axis of the bones, whereas they are perpendicular to this direction in the pseudoteeth. There is no hiatus or evidence of a fusion between the pseudoteeth and the jaw bones. Two possible models for pseudotooth growth are derived from the histological data. The most plausible model is that pseudotooth growth began after the completion of jaw bone growth, as a simple local protraction of periosteal osteogenic activity. Pseudotooth development thus occurred relatively late during ontogeny. The highly vascularized structure and the relative abundance of parallel-fibered bone tissue in the pseudoteeth suggest poor mechanical capabilities. The pseudoteeth were most likely covered and protected by the hardened, keratinized rhamphotheca in the adult during life. The late development of the pseudoteeth would involve a similarly late and/or partial hardening of the rhamphotheca, as displayed by extant Anseriformes, Apterygiformes and some Charadriiformes. This would add support to the hypothesis of a close phylogenetic relationship between Odontopterygiformes and Anseriformes. The late maturation of the Pelagornis feeding apparatus, and hence the delayed capability for efficient prey catching, suggests that Pelagornis was altricial.

Dolichopithecus balcanicus sp. nov., a new Colobinae (Primates, Cercopithecidae) from the early Pliocene of southeastern Europe, with a discussion on the taxonomy of the genus.

We describe a new species of Dolichopithecus, D. balcanicus sp. nov., from the Pliocene (late Ruscinian) of the Balkans. Although known by mandibular remains only, it is readily distinguished from the common European species, D. ruscinensis, by its smaller size, much shallower mandibular corpus, and crowding of the premolars. Ukrainian finds of "Adelopithecus" can be accommodated within D. ruscinensis as a separate, smaller subspecies, D. ruscinensis hypsilophus, but the specimens reported from Asia are more distinct, and they probably represent a separate subgenus of Dolichopithecus, the valid name of which is D. (Parapresbytis) Kalmykov and Mashchenko.

A new hominin from the Basal Member of the Hadar Formation, Dikika, Ethiopia, and its geological context.

In this paper we report for the first time hominin remains from the Basal Member of the Hadar Formation at Dikika, in the Awash Valley of Ethiopia, dating to greater than 3.4 Ma. The new fossil, DIK-2-1, is a fragment of a left mandible and associated dentition. The mandible is attributed to Australopithecus afarensis. However, the new fossil exhibits some metric and morphological features that have not previously been seen in the A. afarensis hypodigm, increasing the already impressive degree of variation in the mandibular sample of the species.

Hominid cranium from Omo: Description and taxonomy of Omo-323-1976-896.

Omo-323-1976-896, a partial hominid cranium dated to ca. 2.1 from the Member G, Unit G-8 of the Shungura Formation, lower Omo Basin of Ethiopia, is described. It is suggested that the specimen is an adult male based on the well-developed and completely fused sagittal crest; heavily worn teeth; relatively large canine; and size of the articular eminence. Omo-323 consists of fragments of the frontal, both temporals, occipital, parietals, and the right maxilla, and is attributed to Australopithecus boisei, making it the oldest known cranium of this species. The specimen shares features with Australopithecus aethiopicus (KNM-WT 17000), thus supporting the existence of an evolving East African robust lineage between ca. 2.6-1.2 Ma. The morphology of Omo-323 increases our knowledge of the intraspecific variability of A.boisei.

A new hominid from the Upper Miocene of Chad, Central Africa.

The search for the earliest fossil evidence of the human lineage has been concentrated in East Africa. Here we report the discovery of six hominid specimens from Chad, central Africa, 2,500 km from the East African Rift Valley. The fossils include a nearly complete cranium and fragmentary lower jaws. The associated fauna suggest the fossils are between 6 and 7 million years old. The fossils display a unique mosaic of primitive and derived characters, and constitute a new genus and species of hominid. The distance from the Rift Valley, and the great antiquity of the fossils, suggest that the earliest members of the hominid clade were more widely distributed than has been thought, and that the divergence between the human and chimpanzee lineages was earlier than indicated by most molecular studies.