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Recent fossil discoveries of early human ancestors from paleoanthropological sites in Africa and elsewhere have demonstrated how various phases of human evolutionary history were much more complica...

Australopitheus anamensis fossils demonstrate that craniodentally and postcranially the taxon was more primitive than its evolutionary successor Australopithecus afarensis. Postcranial evidence suggests habitual bipedality combined with primitive upper limbs and an inferred significant arboreal adaptation. Here we report on A. anamensis fossils from the Assa Issie locality in Ethiopia's Middle Awash area dated to ∼4.2 Ma, constituting the oldest known Australopithecus axial remains. Because the spine is the interface between major body segments, these fossils can be informative on the adaptation, behavior and our evolutionary understanding of A. anamensis. The atlas, or first cervical vertebra (C1), is similar in size to Homo sapiens, with synapomorphies in the articular facets and transverse processes. Absence of a retroglenoid tubercle suggests that, like humans, A. anamensis lacked the atlantoclavicularis muscle, resulting in reduced capacity for climbing relative to the great apes. The retroflexed C2 odontoid process and long C6 spinous process are reciprocates of facial prognathism, a long clivus and retroflexed foramen magnum, rather than indications of locomotor or postural behaviors. The T1 is derived in shape and size as in Homo with an enlarged vertebral body epiphyseal surfaces for mitigating the high-magnitude compressive loads of full-time bipedality. The full costal facet is unlike the extant great ape demifacet pattern and represents the oldest evidence for the derived univertebral pattern in hominins. These fossils augment other lines of evidence in A. anamensis indicating habitual bipedality despite some plesiomorphic vertebral traits related to craniofacial morphology independent of locomotor or postural behaviors (i.e., a long clivus and a retroflexed foramen magnum). Yet in contrast to craniodental lines of evidence, some aspects of vertebral morphology in A. anamensis appear more derived than its descendant A. afarensis.

OBJECTIVES Australopithecus anamensis has comparable δ13 Cenamel values to Ardipithecus ramidus, and both have been characterized as C3 feeders in open woodland habitats similar to "savanna" chimps. Unlike Ar. ramidus and "savanna" chimps, A. anamensis shows a derived dentognathic morphology for tough foods and a dental microwear pattern similar to the C3 -C4 -mixed-feeding A. afarensis. Here I test the hypothesis that changing the variables (e*enamel-diet , δ13 CC3, δ13 CC4 values) used to calculate the percentage of dietary C4 foods (%C4 diet) by 1-2‰ does not make a substantial difference for hominin diet reconstructions [van der Merwe, Masao, & Bamford, 2008, South African Journal of Science 104:153-155]. MATERIALS AND METHODS I estimate vegetation structures for A. anamensis with pedogenic carbonate and faunal enamel δ13 C values from the Pliocene Omo-Turkana Basin (4.2-3.9 Ma). I recalculate A. anamensis' %C4 diet based on new body size-dependent estimates of the e*enamel-diet value and alternative δ13 CC3 and δ13 CC4 values. RESULTS The Pliocene Omo-Turkana Basin shows evidence for dietary resources with a wide range of δ13 C values including canopy-driven 13 C depleted ground C3 foods. Alternative equation variables changed by 1-2‰ yield higher C4 estimates for A. anamensis (15-31%) than previously thought (0-10%). The choice of δ13 CC3 and δ13 CC4 values for estimating %C4 is not a perfunctory task and potentially explains the δ13 C isotopic equifinality of A. anamensis and "savanna" chimps. DISCUSSION My integrative diet model reconciles the carbon isotopic data with the dentognathic and microwear evidence of A. anamensis' diet and suggests that "savanna" chimps are not proper dietary analogs of A. anamensis. A foraging strategy across heterogeneous habitats of the Pliocene Omo-Turkana Basin incorporating an array of 13 C-depleted and 13 C-enriched C3 foods and a portion of C4 resources may have served as one of the selective pressures for A. anamensis, the earliest habitual biped [van der Merwe et al., 2008, South African Journal of Science 104:153-155].

Probabilistic approaches to phylogenetic inference have recently gained traction in paleontological studies. Because they directly model processes of evolutionary change, probabilistic methods facilitate a deeper assessment of variability in evolutionary pattern by weighing evidence for competing models. Although phylogenetic methods used in paleontological studies have generally assumed that evolution proceeds by splitting cladogenesis, extensions to previous models help explore the potential for morphological and temporal data to provide differential support for contrasting modes of evolutionary divergence. Recent methodological developments have integrated ancestral relationships into probabilistic phylogenetic methods. These new approaches rely on parameter-rich models and sophisticated inferential methods, potentially obscuring the respective contributions of data and models. In this study, we describe a simple likelihoodist approach that combines probabilistic models of morphological evolution and fossil preservation to reconstruct both cladogenetic and anagenetic relationships. By applying this approach to a dataset of fossil hominins, we demonstrate the capability of existing models to unveil evidence for anagenesis presented by morphological and temporal data. This evidence was previously recognized by qualitative assessments, but largely ignored by quantitative phylogenetic analyses. For example, we find support for directly ancestral relationships in multiple lineages:Sahelanthropusis ancestral to later hominins;Australopithecus anamensisis ancestral toAu. afarensis;Au. garhiis ancestral toHomo;H. antecessoris ancestral toH. heidelbergensis, which in turn is ancestral to bothH. sapiensandH. neanderthalensis. These results show a benefit of accommodating direct ancestry in phylogenetics. By so doing, quantitative results align more closely with previous qualitative expectations.

Australopithecus anamensis, among the earliest fully bipedal hominin species, lived in eastern Africa around 4 Ma. Much of what is currently known about the paleoecology of A. anamensis comes from the type locality, Kanapoi, Kenya. Here, we extend knowledge of the range of environments occupied by A. anamensis by presenting the first multiproxy paleoecological analysis focusing on Bovidae excavated from another important locality where A. anamensis was recovered, locality 261-1 (ca. 3.97 Ma) at Allia Bay, East Turkana, Kenya. Paleoenvironments are reconstructed using astragalar ecomorphology, mesowear, hypsodonty index, and oxygen and carbon isotopes from dental enamel. We compare our results to those obtained from Kanapoi. Our results show that the bovid community composition is similar between the two fossil assemblages. Allia Bay and Kanapoi bovid astragalar ecomorphology spans the spectrum of modern morphologies indicative of grassland, woodland, and even forest-adapted forms. Dietary reconstructions based on stable isotopes, mesowear, and hypsodonty reveal that these bovids' diet encompassed the full C-3 to C-4 dietary spectrum and overlap in the two data sets. Our results allow us to confidently extend our reconstructions of the paleoenvironments of A. anamensis at Kanapoi to Allia Bay, where this pivotal hominin species is associated with heterogeneous settings including habitats with varying degrees of tree cover, including grasslands, bushlands, and woodlands. (C) 2020 Elsevier Ltd. All rights reserved. Leakey Foundation; Sigma Xi; Explorers Club Washington Group Inc.; Evolving Earth Foundation; Cosmos Club Foundation; Lewis N. Cotlow Fund info:eu-repo/semantics/publishedVersion

A 3.8-million-year-old hominin fossil reveals what the cranium of the oldest known Australopithecus species looked like, casting doubt on assumptions about how these ancient relatives of humans evolved. A fossil reveals what the face of Australopithecus anamensis looked like.

Australopithecus anamensis is a pivotal species in human evolution. It is likely to be the direct ancestor of Australopithecus afarensis and the species that may have given rise to the Homo and Paranthropus lineages. It had a suite of adaptations for habitual bipedalism and a diet that differed from that of earlier hominin species. Under what environmental and ecological conditions did this suite of adaptations arise? The early Pliocene site of Kanapoi in the Lake Turkana Basin of Kenya has the largest sample of A. anamensis in eastern Africa and a rich record of fossil vertebrates. Most Kanapoi fossils are chronologically well constrained by radiometrically dated tephras between the ages of 4.2 and 4.1 million years ago. Sedimentological, isotopic, and faunal data indicate that the environments of Kanapoi during the early Pliocene had a complex range of vegetation types that included closed woodlands, shrubs, and grasslands near a river (for most of the sequence) or lake. These were dynamic landscapes that could shift rapidly from fluvial to lacustrine conditions, and then back. Australopithecus anamensis shared its environments with at least 10 species of very large herbivores, which undoubtedly played a major role in modifying the landscape by opening wooded areas and providing pathways for bipedal hominins. Hominins may have competed for terrestrial resources with abundant suids (Nyanzachoerus and Notochoerus) and for arboreal resources with monkeys (Parapapio being the most common cercopithecid). Kanapoi had a formidable group of predators that included a very abundant species of hyena (Parahyaena howelli), two sabre-tooth felids (Dinofelis and Homotherium), a giant otter (Enhydriodon cf. dikikae), and three species of crocodiles. Various measures of abundance indicate that A. anamensis was an important component of the Kanapoi early Pliocene ecosystems, and that its key adaptations allowed this species to thrive in complex and dynamic landscapes. (C) 2019 Elsevier Ltd. All rights reserved. Leakey Foundation Paleontological Scientific Trust (PAST) of South Africa National Science FoundationNational Science Foundation (NSF) University of Missouri Research Board info:eu-repo/semantics/publishedVersion

Enamel thickness remains an important morphological character in hominin systematics and is regularly incorporated into dietary reconstructions in hominin species. We expand upon a previous study of enamel thickness in mandibular molars by examining a large maxillary molar sample of Plio-Pleistocene hominins (n = 62) and a comparative sample of extant nonhuman apes (n = 48) and modern humans (n = 29). 2D mesial planes of section were generated through microtomography, and standard dental tissue variables were measured to calculate average enamel thickness (AET) and relative enamel thickness (RET). AET was also examined across the lingual, occlusal, and buccal regions of the crown. This study confirms previous findings of increasing enamel thickness throughout the Plio-Pleistocene, being thinnest in Australopithecus anamensis and peaking in Australopithecus boisei, with early Homo specimens, exhibiting intermediate enamel thickness. Agreeing with previous findings, 2D plane of section enamel thickness is found to be a poor taxonomic discriminator, with no statistically significant differences observed between fossil hominins. For fossil hominins, modern humans, and Pongo, the occlusal region of enamel was the thickest, and the lingual enamel thickness was greater than buccal. Pan and Gorilla present the opposite pattern with enamel being thinnest occlusally. Comparison at each molar position between the maxilla and mandible revealed very few significant differences in fossil hominins but some evidence of significantly thicker maxillary enamel (AET) in modern humans and thinner maxillary enamel in Pan (RET).

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.

Woranso-Mille is a paleoanthropological site in Ethiopia sampling an important and under-represented time period in human evolution (3.8–3.6 million years ago). Specimens of cf. Australopithecus anamensis, Australopithecus afarensis, and the recently named Australopithecus deyiremeda have been recovered from this site. Using multiple habitat proxies, this study provides a paleoecological reconstruction of two fossiliferous collection areas from Woranso-Mille, Aralee Issie (ARI) and Mesgid Dora (MSD). Previous reconstructions based on faunal assemblages have pointed, due to the presence of aepycerotins, alcelaphins, and proboscideans, to the existence of open habitats as well as more closed ones, based on the occurrence of cercopithecids, giraffids, and traglephins. Results from community structure analysis (proportions of locomotor and dietary adaptations) at ARI and MSD indicated a predominance of open habitats, such as shrublands. Mesowear analysis revealed that ungulates of all dietary types (grazers, leaf and fruit browsers, and mixed feeders) were present in nearly equal proportions. Ecomorphological analyses using linear measurements of the astragalus and phalanges indicated that bovids utilizing locomotor behaviors associated with all habitat types were present, though the intermediate-cover habitat bovids were best represented in the sample (Heavy cover at ARI and Light cover at MSD). Together, these results suggest that the ARI and MSD localities were heterogeneous habitats (mosaics), likely with densely vegetated areas along a paleo-river and more open regions (woodlands, grasslands) available away from the river.

Stable isotope analysis has long been used to study the dietary ecology of living and fossil primates, and there has been increasing interest in using stable isotopes to study primate habitat use and anthropogenic impacts on non-human primates. Here, we examine the stable carbon and nitrogen isotope compositions of chimpanzees (Pan troglodytes) from seven communities in Uganda across a continuum of habitat structure (closed to more open) and access to anthropogenic resources (no reliance to heavy reliance). In general, the hair δ(13) C, but not δ(15) N, values of these communities vary depending on forest structure and degree of anthropogenic influence. When integrated with previously published hair δ(13) C and δ(15) N values for Pan, it is apparent that modern "savanna" and "forest" Pan form discrete clusters in carbon and nitrogen isotope space, although there are exceptions probably relating to microhabitat specialization. The combined dataset also reveals that Pan δ(13) C values (but not δ(15) N values) are inversely related to rainfall (r(2) = 0.62). We converted Pan hair δ(13) C values to enamel equivalents and made comparisons to the fossil hominoids Sivapithecus sp., Gigantopithecus blacki, Ardipithecus ramidus, and Australopithecus anamensis. The δ(13) C values of the fossil hominins Ar. ramidus and Au. anamensis do not cluster with the δ(13) C values of modern Pan in "forest" habitats, or with fossil hominoids that are believed to have inhabited forests. Am. J. Primatol. 78:1070-1085, 2016. © 2016 Wiley Periodicals, Inc.

The mandibular third premolar (P3) exhibits substantial differences in size and shape among hominoid taxa, and displays a number of discrete traits that have proven to be useful in studies of hominin taxonomy and phylogeny. Discrete traits at the enamel-dentine junction (EDJ) can be accurately assessed on moderately worn specimens, and often appear sharper than at the outer-enamel surface (OES). Here we use microtomography to image the P3 EDJ of a broad sample of extant apes, extinct hominins and modern humans (n = 100). We present typologies for three important premolar discrete traits at the EDJ (transverse crest, marginal ridge and buccal grooves), and score trait frequencies within our sample. We find that the transverse crest is variable in extant apes, while the majority of hominins display a transverse crest which runs directly between the two major premolar cusps. Some Neanderthals display a unique form in which the transverse crest fails to reach the protoconid. We find that mesial marginal ridge discontinuity is common in Australopithecus anamensis and Australopithecus afarensis while continuous marginal ridges largely characterize Australopithecus africanus and Paranthropus. Interrupted mesial and distal marginal ridges are again seen in Homo sapiens and Neanderthals. Premolar buccal grooves, previously identified at the OES as important for hominin systematics, are again found to show a number of taxon-specific patterns at the EDJ, including a clear difference between Australopithecus and Paranthropus specimens. However, their appearance may be dependent on the morphology of other parts of the crown such as the protoconid crest, and the presence of accessory dentine horns. Finally, we discuss rare variations in the form of dentine horns that underlie premolar cusps, and their potential homology to similar morphologies in other tooth positions.

The early hominin (Ardipithecus and Australopithecus) fossil record contains over 100 preserved vertebral elements (n = 107; approximately half of which are well-preserved), ~65% of which have not been described since the turn of the millennium. Many are fragments, some for which detailed descriptions are pending (e.g., those of Australopithecus anamensis). Australopithecus afarensis and Australopithecus sediba are known from cervical, thoracic, and lumbar vertebrae, whereas Australopithecus africanus is known from thoracic and lumbar vertebrae but not cervical vertebrae. A partial skeleton from Member 4 of Sterkfontein, StW 573, preserves vertebrae from all presacral regions, but its species designation is debated and not yet formalized in the literature. Other early hominin species, such as Sahelanthropus tchadensis, Orrorin tugenensis, Ardipithecus kadabba, Australopithecus deyiremeda, Australopithecus bahrelghazali, and Australopithecus garhi, do not preserve vertebrae. Vertebrae from Swartkrans and Cooper’s Cave are thought to belong to either Paranthropus or Homo and are discussed in Meyer and Williams (this volume). The vertebrae discussed in this chapter are from five sites in East and South Africa: Aramis, Asa Issie, and Hadar from the Afar Depression of Ethiopia and Sterkfontein and Malapa in the Cradle of Humankind, South Africa.

Carbon isotope ratios of mammalian teeth from the Kanapoi site in northern Kenya are interpreted in the context of C3 and C4 derived resources to investigate the paleoecology of Australopithecus anamensis. δ13C values of large mammals, when compared at the taxon level, show an ecosystem that is strongly biased towards mixed feeders and browsers. However, sufficient C4 resources were present such that some C4 dominated grazers were also present in the large mammal fauna. Analyses of micromammals shows that their diets were C3 dominated or C3-C4 mixed. Carbon isotope studies of primates shows that the major primate tribes-Colobini, Papioini, Hominini-all made some use of C4 resources in their respective diets; the Hominini had a higher fraction of C3 diet resources than the other primate tribes represented in the fossil record.

Reconstructed habitats of Australopithecus anamensis at Kanapoi by Wynn (2000) yielded evidence for both wooded and grassy environments. Wynn's study was based on stable isotopic (δ13CPC, δ18OPC) analyses of a small sample of pedogenic nodules (n = 14) collected from paleosols spanning Kanapoi's stratigraphic interval. Whether this small sample size adequately characterized Kanapoi's vegetation or was the result of time averaging remains unclear. To address this uncertainty, we sampled Kanapoi paleosols at 39 locations (78 analyses) from laterally extensive units. Our data demonstrate that Kanapoi offered A. anamensis diverse habitats distributed in temporally discrete stratigraphic horizons. Habitat heterogeneity appears to have been a real aspect of Kanapoi paleoenvironments and not an artifact of Wynn's (2000) small sample size or time averaging. We suggest habitat heterogeneity was influenced by the location of Kanapoi at the confluence of fluvial, deltaic, and lacustrine depositional environments. We also compared Kanapoi's δ13CPC and δ18OPC values to those of other Pliocene hominin localities in eastern Africa dated to 4.5–3.7 Ma. Kanapoi's δ18OPC values are significantly higher than most sites, potentially reflecting regional variability in water source δ18O values and/or more arid climatic conditions. Kanapoi's δ13CPC values indicate significantly more woody cover than at all other sites except those in the Turkana Basin. Kanapoi provided A. anamensis with a wide range of C3–C4 resources as the C4 biome spread across eastern Africa. [Wynn, J.G., 2000. Paleosols, stable carbon isotopes and paleoenvironmental interpretation of Kanapoi, Northern Kenya. J. Hum. Evol. 39, 411–432.]

Fossil bird remains from the Pliocene hominin-bearing locality of Kanapoi comprise >100 elements representing at least 10 avian families, including previously undescribed elements referred to the ‘giant’ Pliocene marabou stork Leptoptilos cf. falconeri. The taxonomic composition of the Kanapoi fossil avifauna reveals an assemblage with a substantial aquatic component, corroborating geological evidence of this locality's close proximity to a large, slow-moving body of water. Both the taxonomic composition and relative abundance of avian higher-level clades at Kanapoi stand in stark contrast to the avifauna from the slightly older (∼4.4 Ma vs. 4.2 Ma) hominin-bearing Lower Aramis Member of Ethiopia, which has been interpreted as representing a mesic woodland paleoenvironment far from water. In general, the taxonomic composition of the Kanapoi avifauna resembles that from the Miocene hominoid-bearing locality of Lothagam (though Kanapoi is more diverse), and the aquatic character of the Kanapoi avifauna supports the idea that the environmental conditions experienced by Australopithecus anamensis at Kanapoi were markedly different from those experienced by Ardipithecus ramidus at Aramis. Additionally, the relative abundance of marabou stork (Leptoptilos) remains at Kanapoi may suggest a longstanding commensal relationship between total-clade humans and facultatively scavenging marabous. Additional avian remains from nearby fossil localities (e.g., the Nachukui Formation), ranging in age from 3.26 to 0.8 Ma, reveal the long-term persistence of an aquatic avifauna in the region.

Reliable estimates of when hominin taxa originated and went extinct are central to addressing many paleoanthropological questions, including those relating to macroevolutionary patterns. The timing of hominin temporal ranges can be used to test chronological predictions generated from phylogenetic hypotheses. For example, hypotheses of phyletic ancestor–descendant relationships, based on morphological data, predict no temporal range overlap between the two taxa. However, a fossil taxon's observed temporal range is almost certainly underestimated due to the incompleteness of both the fossil record itself and its sampling, and this decreases the likelihood of observing temporal overlap. Here, we focus on a well-known and widely accepted early hominin lineage, Australopithecus anamensis–afarensis, and place 95% confidence intervals (CIs) on its origination and extinction dates. We do so to assess whether its temporal range is consistent with it being a phyletic descendant of Ardipithecus ramidus and/or a direct ancestor to the earliest claimed representative of Homo (i.e., Ledi-Geraru). We find that the last appearance of Ar. ramidus falls within the origination CI of Au. anamensis–afarensis, whereas the claimed first appearance of Homo postdates the extinction CI. These results are consistent with Homo evolving from Au. anamensis–afarensis, but temporal overlap between Ar. ramidus and Au. anamensis–afarensis cannot be rejected at this time. Though additional samples are needed, future research should extend our initial analyses to incorporate the uncertainties surrounding the range endpoints of Ar. ramidus and earliest Homo. Overall, our findings demonstrate the need for quantifying the uncertainty surrounding the appearances and disappearances of hominin taxa in order to better understand the timing of evolutionary events in our clade's history.

Stable carbon isotope studies suggest that early hominins may have diversified their diet as early as 3.76 Ma. Early Pliocene hominins, including Australopithecus anamensis, had diets that were dominated by C3 resources while Late Pliocene hominins, including Australopithecus afarensis—a putative descendant of A. anamensis—had diets that included both C3 and C4 resources. It has been hypothesized that the expansion of C4 grasslands in eastern Africa during the Pliocene could have prompted hominins to incorporate C4 resources in their diets. However, dental microwear analyses suggest that diet diversification did not involve changes in the mechanical properties of the foods consumed. To provide contextual and comparative information on this issue, the diet of suids from the A. anamensis site of Kanapoi and the A. afarensis site of Hadar is investigated. Using dental microwear texture analyses, it is shown that despite significant dietary overlap, there is evidence for dietary niche partitioning among suids. Based on comparisons with the diet of extant African suids, it is inferred that Nyanzachoerus pattersoni (n = 21) was a mixed feeder, Nyanzachoerus jaegeri (n = 4) and Notochoerus euilus (n = 61) were habitual grazers, and Kolpochoerus afarensis (n = 34) had a broad diet that included hard brittle foods and underground resources. The dental microwear of Ny. pattersoni and Ny. jaegeri/No. euilus do not differ significantly between Kanapoi and Hadar. Most differences are driven by K. afarensis, a suid absent at Kanapoi but present at Hadar. Food availability probably differed between Hadar and Kanapoi, and it is likely that A. afarensis did not exploit some of the foods (e.g., underground resources) consumed by suids. It is hypothesized that despite the incorporation of C4 resources in the diet, a significant dietary change towards flexible diets in the hominin lineage had yet to come.

Here we present the first full anatomical description of the 3.67 million-year-old Australopithecus skull StW 573 that was recovered with its skeleton from the Sterkfontein Member 2 breccia in the Silberberg Grotto. Analysis demonstrates that it is most similar in multiple key morphological characters to a group of fossils from Sterkfontein Member 4 and Makapansgat that are here distinguished taxonomically as Australopithecus prometheus. This taxon contrasts with another group of fossils from those sites assigned to Australopithecus africanus. The anatomical reasons for why these groupings should not be lumped together (as is frequently done for the South African fossils) are discussed in detail. In support of this taxonomy, we also present for the first time a newly reconstructed palate of A. africanus (StW 576 from Sterkfontein Member 4), which has a uniquely complete permanent dentition. The StW 573 skull also has certain similarities with other earlier Australopithecus fossils in East Africa, assigned to Australopithecus afarensis and Australopithecus anamensis, which are discussed. One of its most interesting features is a pattern of very heavy anterior dental wear unlike that found in A. africanus but resembling that found in A. anamensis at 4.17 Ma. Because the StW 573 skull is associated with a near-complete skeleton that is also described for the first time in this special issue, we are now able to use this individual to improve our understanding of more fragmentary finds in the South African fossil record of Australopithecus.

Recent fieldwork at Kanapoi has expanded the sample of fossil cercopithecids, facilitating a re-appraisal of their taxonomy. The assemblage now includes at least one species of cercopithecin, two papionins, and two colobines. The guenon Nanopithecus browni is similar in dental size to extant Miopithecus. We tentatively re-affirm the identification of Parapapio cf. ado and confirm the presence of Theropithecus. The colobines include a small form tentatively attributed to Kuseracolobus and a second larger species. The Kanapoi fossils represent the oldest occurrences of guenons in Africa and of the important genus Theropithecus, the most abundant and widespread primate in the Neogene of Africa. In the assemblage, Parapapio cf. ado is the most abundant form, comprising the majority of specimens. All of the other taxa are comparatively rare. Colobines make up a small part of the Kanapoi fossil assemblage compared to most other contemporary sites, including Allia Bay, Kenya, where, like Kanapoi, Australopithecus anamensis has been found. The presence of Theropithecus is consistent with the presence of some relatively open habitat at Kanapoi. While the ecological preferences of the small cercopithecin are unknown, most guenons are associated with relatively wooded habitats, as are most colobines, suggesting the availability of at least some wooded areas.

The study considers the turnover in hominins, together with carnivorans and other primates, at 3.5 Ma against an environmental backdrop. Communalities are identified between evolving guilds that may directly inform hominin evolution. These are the evolution of (a) dietary generalists and (b) evidence for sociality in carnivores, baboons and hominins. Sociality and behavioural flexibility are regarded advantageous for the procurement of resources while, at the same time, reducing intraspecific competition; in primates it may initially also have served to reduce predation risk. Behavioural flexibility explains the evolutionary success of Panthera leo, Papio and Homo. Viewed within a wider palaeoecological and environmental context, it is possible that sociality in hominins, including allocare, were triggered by abiotic changes at about 3.5 Ma. If confirmed in future studies, this would mark the beginning of hominin life history evolution.

In a find that some are calling one of the most spectacular hominin discoveries of the decade, fossil hunters working in northeastern Ethiopia have uncovered a nearly complete, 3.8-million-year-old cranium belonging to the oldest known species of australopiths, Australopithecus anamensis. The ancient hominin, long thought to be the direct precursor to the "Lucy" species, A. afarensis, was previously known mostly from jawbones and teeth. Most researchers tend to think A. anamensis gradually transitioned into and was replaced by A. afarensis. But in a pair of studies describing and interpreting the new skull, paleoanthropologists argue there9s now evidence the two australopith species actually overlapped for about 100,000 years. Other researchers caution that many more fossils will need to be found before they accept that version of hominin history.

Excavations at Kanapoi in north-western Kenya have yielded the most numerically abundant and taxonomically diverse early Pliocene (4.19 Ma) terrestrial small mammal assemblage known from Kenya. A minimum of 15 species are reported, including soricids, sengis, leporids, and rodents: all taxa are referable to extant genera, with the exception of the murine rodent, Saidomys. The majority of the terrestrial small mammals are derived from a bone bed at Nzube's Mandible Site, closely associated with the holotype mandible of Australopithecus anamensis. A smaller number were surface-collected or obtained from screening at several other sites, including the Bat Site. Most small mammals from Nzube's Mandible Site and the Bat Site likely represent prey accumulated as regurgitated pellets from owls, in particular barn owls or giant eagle owls. The small mammal fauna is dominated by the spiny mouse, Acomys: the next most commonly recovered taxa are the multimammate mouse, Mastomys, and the African gerbil, Gerbilliscus. Comparisons of the Kanapoi fauna to other eastern African late Miocene-Pliocene (and one Pleistocene) faunas at the generic level suggest the greatest similarity is to Lemudong'o, Kenya, and Omo B and Aramis, Ethiopia. Further similarities with other localities such as Laetoli, Tanzania, and Hadar, Ethiopia, suggest the existence of a corridor for dispersal along the East African Rift Valley between Ethiopia, Kenya, and Tanzania in the early Pliocene. Further comparisons of the relative abundances of individuals in different families (or subfamilies) emphasize the distinctiveness of the Kanapoi small mammal fauna. The Kanapoi fauna is likely derived from a heterogeneous but relatively arid environment.

Isolated amphibian bones from the early Pliocene of Kanapoi (West Turkana, Kenya) help to improve the scarce fossil record of the late Neogene and Quaternary amphibians from East Africa. All currently available 579 bones are referable exclusively to the Anura (frogs and toads). More than half of the remains (366) are identified as Hemisus cf. Hemisus marmoratus, an extant species that still inhabits Kenya, but apparently not the northwest of the country and the Turkana area in particular. The rest of the remains are identified simply as Anura indet. because of poor preservation or non congruence with the relatively few African extant taxa whose osteology is known in detail. The Hemisus material represents the first fossil record for Hemisotidae, an endemic African family of peculiar, head-first burrowing frogs, whose sister taxon relationships indicate a divergence from brevicipitids in the Late Cretaceous or early Paleocene. The ecological requirements of extant H. marmoratus suggest that the Kanapoi area surrounding the fluvial and deltaic settings, from where the fossil remains of vertebrates were buried, was likely a grassland or relatively dry, open low tree-shrub savanna.

Kanapoi, Kenya, has yielded the earliest evidence of the genus Australopithecus , Australopithecus anamensis . Renewed fieldwork from 2012 through 2015 yielded 18 new fossils attributable to this species. The new specimens include the second maxillary fragment known from a Kanapoi hominin and the first from a relatively young adult. The new maxilla has the distinctive rounded nasal aperture margin characteristic of A. anamensis . A second partial proximal tibia from the site is the first postcranial element from a small A. anamensis individual. A new partial mandible and complete mandibular dentition display distinctive Kanapoi hominin morphology, but the mandible displays a larger trigonid on its fourth premolar than any known so far. Two new complete sets of mandibular incisors are also notably large, especially the lateral ones, a distinctive feature of A. anamensis compared with Australopithecus afarensis . The new fossils also highlight the distinctive morphology of Kanapoi A. anamensis compared to later hominins.

Reconstructions of habitat at sites like Kanapoi are key to understanding the environmental circumstances in which hominins evolved during the early Pliocene. While Australopithecus anamensis shows evidence of terrestrial bipedality traditionally associated with a more open setting, its enamel has low δ13C values consistent with consumption of C3 foods, which predominate in wooded areas of tropical Africa. Habitat proxies, ranging from paleosols and their carbonates to associated herbivore fauna and their carbon isotope ratios, suggest a heterogeneous setting with both grass and woody plant components, though the proportions of each have been difficult to pin down. Here we bring dental microwear texture analysis of herbivorous fauna to bear on the issue. We present texture data for fossil bovids, primates, rodents, and suids (n = 107 individuals in total) from the hominin bearing deposits at Kanapoi, and interpret these in the light of closely related extant mammals with known differences in diet. The Kanapoi bovid results, for example, are similar to those for extant variable grazers or graze-browse intermediate taxa. The Kanapoi suid data vary by taxon, with one similar to the pattern of extant grazers and the other more closely resembling mixed feeders. The Kanapoi primates and rodents are more difficult to associate with a specific environment, though it seems that grass was likely a component in the diets of both. All taxa evince microwear texture patterns consistent with a mosaic of discrete microhabitats or a heterogeneous setting including both tree and grass components.

The Pliocene hominins Australopithecus anamensis and Australopithecus afarensis likely represent ancestor-descendent taxa--possibly an anagenetic lineage--and capture significant change in the morphology of the canine and mandibular third premolar (P(3)) crowns, dental elements that form the canine honing complex in nonhuman catarrhines. This study focuses on the P(3) crown, highlighting plesiomorphic features in A. anamensis. The A. afarensis P(3) crown, in contrast, is variable in its expression of apomorphic features that are characteristic of geologically younger hominins. Temporal variation characterizes each taxon as well. The A. anamensis P(3) from Allia Bay, Kenya expresses apomorphic character states, shared with A. afarensis, which are not seen in the older sample of A. anamensis P(3)s from Kanapoi, Kenya, while spatiotemporal differences in shape exist within the A. afarensis hypodigm. The accumulation of derived features in A. afarensis results in an increased level of P(3) molarisation. P(3) molarisation did not evolve concurrent with postcanine megadontia and neither did the appearance of derived aspects of P(3) occlusal form coincide with the loss of canine honing in hominins, which is apparent prior to the origin of the genus Australopithecus. A. afarensis P(3) variation reveals the independence of shape, size, and occlusal form. The evolution of the P(3) crown in early Australopithecus bridges the wide morphological gap that exists between geologically younger hominins on the one hand and extant apes and Ardipithecus on the other.

Many researchers have suggested that Australopithecus anamensis and Australopithecus afarensis were among the earliest hominins to have diets that included hard, brittle items. Here we examine dental microwear textures of these hominins for evidence of this. The molars of three Au. anamensis and 19 Au. afarensis specimens examined preserve unobscured antemortem microwear. Microwear textures of these individuals closely resemble those of Paranthropus boisei , having lower complexity values than Australopithecus africanus and especially Paranthropus robustus . The microwear texture complexity values for Au. anamensis and Au. afarensis are similar to those of the grass-eating Theropithecus gelada and folivorous Alouatta palliata and Trachypithecus cristatus . This implies that these Au. anamensis and Au. afarensis individuals did not have diets dominated by hard, brittle foods shortly before their deaths. On the other hand, microwear texture anisotropy values for these taxa are lower on average than those of Theropithecus , Alouatta or Trachypithecus . This suggests that the fossil taxa did not have diets dominated by tough foods either, or if they did that directions of tooth–tooth movement were less constrained than in higher cusped and sharper crested extant primate grass eaters and folivores.

The earliest evidence ofAustralopithecusgoes back toca4.2 Ma with the first recorded appearance ofAustralopithecus ‘anamensis’at Kanapoi, Kenya.Australopithecus afarensisis well documented between 3.6 and 3.0 Ma mainly from deposits at Laetoli (Tanzania) and Hadar (Ethiopia). The phylogenetic relationship of these two ‘species’ is hypothesized as ancestor–descendant. However, the lack of fossil evidence from the time between 3.6 and 3.9 Ma has been one of its weakest points. Recent fieldwork in the Woranso-Mille study area in the Afar region of Ethiopia has yielded fossil hominids dated between 3.6 and 3.8 Ma. These new fossils play a significant role in testing the proposed relationship betweenAu. anamensisandAu. afarensis. The Woranso-Mille hominids (3.6–3.8 Ma) show a mosaic of primitive, predominantlyAu. anamensis-like, and some derived (Au. afarensis-like) dentognathic features. Furthermore, they show that, as currently known, there are no discrete and functionally significant anatomical differences betweenAu. anamensisandAu. afarensis. Based on the currently available evidence, it appears that there is no compelling evidence to falsify the hypothesis of ‘chronospecies pair’ or ancestor–descendant relationship betweenAu. anamensisandAu. afarensis. Most importantly, however, the temporally and morphologically intermediate Woranso-Mille hominids indicate that the species namesAu. afarensisandAu. anamensisdo not refer to two real species, but rather to earlier and later representatives of a single phyletically evolving lineage. However, if retaining these two names is necessary for communication purposes, the Woranso-Mille hominids are best referred to asAu. anamensisbased on new dentognathic evidence.

The Middle Awash Ardipithecus ramidus sample comprises over 145 teeth, including associated maxillary and mandibular sets. These help reveal the earliest stages of human evolution. Ar. ramidus lacks the postcanine megadontia of Australopithecus . Its molars have thinner enamel and are functionally less durable than those of Australopithecus but lack the derived Pan pattern of thin occlusal enamel associated with ripe-fruit frugivory. The Ar. ramidus dental morphology and wear pattern are consistent with a partially terrestrial, omnivorous/frugivorous niche. Analyses show that the ARA-VP-6/500 skeleton is female and that Ar. ramidus was nearly monomorphic in canine size and shape. The canine/lower third premolar complex indicates a reduction of canine size and honing capacity early in hominid evolution, possibly driven by selection targeted on the male upper canine.

Hominid fossils predating the emergence of Australopithecus have been sparse and fragmentary. The evolution of our lineage after the last common ancestor we shared with chimpanzees has therefore remained unclear. Ardipithecus ramidus , recovered in ecologically and temporally resolved contexts in Ethiopia’s Afar Rift, now illuminates earlier hominid paleobiology and aspects of extant African ape evolution. More than 110 specimens recovered from 4.4-million-year-old sediments include a partial skeleton with much of the skull, hands, feet, limbs, and pelvis. This hominid combined arboreal palmigrade clambering and careful climbing with a form of terrestrial bipedality more primitive than that of Australopithecus . Ar. ramidus had a reduced canine/premolar complex and a little-derived cranial morphology and consumed a predominantly C 3 plant–based diet (plants using the C 3 photosynthetic pathway). Its ecological habitat appears to have been largely woodland-focused. Ar. ramidus lacks any characters typical of suspension, vertical climbing, or knuckle-walking. Ar. ramidus indicates that despite the genetic similarities of living humans and chimpanzees, the ancestor we last shared probably differed substantially from any extant African ape. Hominids and extant African apes have each become highly specialized through very different evolutionary pathways. This evidence also illuminates the origins of orthogrady, bipedality, ecology, diet, and social behavior in earliest Hominidae and helps to define the basal hominid adaptation, thereby accentuating the derived nature of Australopithecus .

The Woranso-Mille paleontological study area is located in the central Afar region of Ethiopia, about 360 km northeast of the capital, Addis Ababa. Some parts of this area have been paleontologically known since the 1970s. However, most of the fossiliferous areas were discovered by surveys conducted in the region between 2002 and 2004. By the end of the 2006 field season, a total of 17 vertebrate localities had been designated, and more than 1000 vertebrate fossil specimens collected. Among these specimens, there are more than 20 fossil hominid craniodental and postcranial remains, including one partial skeleton, of Pliocene age (3.5–3.8 Ma). Research at this study area has thus far focused on the geology and paleontology of the early Pliocene deposits along the Mille river and adjacent areas located between the towns of Mille and Chifra. Exposed sediments in the new fossiliferous area are mostly silty sand and silty clay horizons interbedded with a number of volcanic tuffs and basaltic flows suitable for 40Ar/39Ar radioisotopic dating. The total stratigraphic section is about 50 m thick with a minimum age of ~3.5 Ma. The study area also has deposits of early to middle Pleistocene age, although no locality has been designated to date. The new Woranso-Mille paleontological study area provides a crucial temporal window into the time during which Australopithecus anamensis (3.9–4.2 Ma) appears to have given rise to Australopithecus afarensis (3.0–3.6 Ma). Radiometric dates have thus far yielded a minimum age of 3.5 Ma for the hominid localities and this conforms well with a biochronological age estimate of 3.6–3.8 Ma. The associated fauna, particularly the abundance of fossil cercopithecids and presence of diverse aquatic fossil vertebrates, indicates a relatively closed, wooded habitat probably associated with a paleo-Mille river.

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 β- 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.

Molar microwear fabrics in extant mammals vary with diet and, more particularly, the physical properties of the items that are consumed. Praeanthropus afarensis is well represented in the fossil record over a prolonged and radiometrically controlled temporal span, and reasonably robust paleoecological reconstructions are available for the various localities from which it is known. We therefore examined molar microwear in this species to determine whether diet varied in relation to time or in response to different ecological conditions. Of more than 70 specimens of Pr. afarensis that contain one or more worn permanent molars, only 19 were found to be suitable for microwear analysis. These derive from eight temporal horizons in the Laetolil Beds and Hadar Formation spanning approximately 400 kyr (3.6–3.2 Ma). Six paleoecological categories have been reconstructed for these horizons, and these were ranked on the basis of floral cover. None of the microwear variables observed for Pr. afarensis is significantly associated with either temporal or paleoecological rank. Thus, microwear and, by extension, diet does not appear to have altered significantly in Pr. afarensis through time or in response to different paleoecological circumstances. The wear pattern that appears to have characterized Pr. afarensis overlaps extensively that of Gorilla gorilla beringei and differs notably from the fabrics of extant primates (e.g., Cebus apella and Cercocebus albigena ) that consume hard objects. The high proportion of scratches on Pr. afarensis molars suggests the inclusion of fine abrasives in or on the food items consumed by those individuals sampled in this study. Although Pr. afarensis may have been morphologically equipped to process hard, brittle items, the microwear data suggest that it did not necessarily do so, even in the face of varying environmental circumstances. Explanatory scenarios that describe Pr. afarensis as part of an evolutionary trajectory involving a more heavily masticated diet with an increased reliance on hard, brittle items need to be reconsidered. However, fallback foods that were consumed during relatively short, albeit critical periods may have exerted sufficient selective pressure to explain the evolution of the comparatively robust Pr. afarensis trophic apparatus. Because it is unlikely that many individuals from such restricted temporal intervals would be sampled in the paleontological record, we suggest that the most productive approach to the elucidation of paleodiet is the integration of genetic (morphological) and epigenetic (microwear and isotopic) lines of evidence.

The discovery of 4.5-million-year-old fossils of the hominid Ardipithecus ramidus increases our knowledge of a fascinating stage of human evolution, before hominids left the forests for the open savanna. Fossil finds of this age are rare, but deposits in Gona, Ethiopia, have yielded material from at least nine individuals. Their context shows that they lived in an environment of moderate rainfall woodland, and grasslands. Comparative biomolecular studies suggest that the last common ancestor of humans and chimpanzees, our closest living relatives, lived during the Late Miocene–Early Pliocene1,2. Fossil evidence of Late Miocene–Early Pliocene hominid evolution is rare and limited to a few sites in Ethiopia3,4,5, Kenya6 and Chad7. 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.

In an earlier study, stress lines in primate teeth were found to occur on a recurrent basis, probably corresponding to seasonal fluctuations in environmental parameters, such as food availability (Macho et al., J. Hum. Evol. 30 (1996) 57–70). In the present study this approach was extended to the study of teeth of extant and extinct mammals, with the specific aim to determine the pattern of seasonality at the Australopithecus anamensis -bearing site at Allia Bay, Kenya. It was found that extant and extinct species, who share similar dietary/ecological adaptations, are comparable in their patterns of stress. Typical browsers/mixed feeders were found to exhibit three recurrent disturbances per year, whereas grazers usually only exhibit two. The average spacing between lines is also comparable between extant and extinct species. Hence, while the severity and predictability of the seasons probably fluctuated during crucial periods of hominin evolution, there is little doubt that all hominins lived in a seasonal environment. At Allia Bay, the pattern of stress lines found in mammals suggests that the environmental conditions in which A. anamensis lived may have been comparable to those found in the Masai Mara today.

Carbon ( 13 C: 12 C) and oxygen ( 18 O: 16 O) stable isotope ratio analysis was performed on well-preserved tooth enamel carbonate from fossil fauna recovered from a single excavation at the early hominid site of Allia Bay, East Turkana, Kenya. These data show greater enrichment in both 12 C and in 16 O than expected, based on the oxygen isotope composition of the middle Pliocene ocean, and on todays ecology. The pattern of these data argues against a diagenetic explanation for the enrichment. The carbon stable isotope data of known browsers suggest a more extensive canopy cover during the middle Pliocene than todays environments. The presence of browsing pig genera, hippo genera, deinotheres, and giraffes with d 13 C values more negative than todays all argue for woodland habitats. The presence of several grazing genera point to the presence of grasslands as well. The oxygen stable isotope ratios indicate that the site was better-watered than today, although the source, seasonal pattern, and actual amount of water cannot be determined from these data. The overall mosaic of environments suggested by these data, in combination with reports of exotic trees recovered in nearby deposits, indicate that woodlands were present in the region 3.9 my, unlike today. Such a setting matches expectations for the selective advantages of nut-eating, bipedal hominids over other hominoids. 2003 Elsevier Inc. All rights reserved.

Cosmogenic aluminum-26 and beryllium-10 burial dates of low-lying fossiliferous breccia in the caves at Sterkfontein, South Africa, show that associated hominid fossils accumulated in the Lower Pliocene. These dates indicate that the skeleton StW 573 and newly discovered specimens from Jacovec Cavern have much the same age: approximately 4 million years. These specimens are thus of an age similar to Australopithecus anamensis from East Africa.

Received 18 March 2014, Accepted 27 February 2015, Available online 24 March 2015, A new collection of 49,000 year old Neandertal fossil humeri from the El Sidrón cave site (Asturias, Spain) is presented. A total of 49 humeral remains were recovered, representing 10 left and 8 right humeri from adults, adolescents, and a juvenile (not included in the analyses). 3D geometric morphometric (GM) methods as well as classic anthropological variables were employed to conduct a broad comparative analysis by means of mean centroid size and shape comparisons, principal components analysis, and cluster studies. Due to the fragmentary nature of the fossils, comparisons were organized in independent analyses according to different humeral portions: distal epiphysis, diaphysis, proximal epiphysis, and the complete humerus. From a multivariate viewpoint, 3D-GM analyses revealed major differences among taxonomic groups, supporting the value of the humerus in systematic classification. Notably, the Australopithecus anamensis (KP-271) and Homo ergaster Nariokotome (KNM-WT 15000) distal humerus consistently clusters close to those of modern humans, which may imply a primitive condition for Homo sapiens morphology. Australopithecus specimens show a high degree of dispersion in the morphospace. The El Sidrón sample perfectly fits into the classic Neandertal pattern, previously described as having a relatively wide olecranon fossa, as well as thin lateral and medial distodorsal pillars. These characteristics were also typical of the Sima de los Huesos (Atapuerca) sample, African mid-Pleistocene Bodo specimen, and Lower Pleistocene TD6-Atapuerca remains and may be considered as a derived state. Finally, we hypothesize that most of the features thought to be different between Neandertals and modern humans might be associated with structural differences in the pectoral girdle and shoulder joint., Ministerio de Economía y Competitividad of Spain: CGL2012-36682 and CGL2012-37279., Convenio Principado de Asturias-Universidad de Oviedo CN-09-084.

Remains of earlyHomo andParanthropus have been recovered from two contemporaneous sites (Uraha and Malema) in the “Hominid Corridor” in Northern Malawi (Chiwondo Beds). Faunal dating suggests an age of 2.5–2.3 Ma for both hominids. The two specimens, a mandible attributed toHomo rudolfensis (UR 501 from Uraha), and a maxillary fragment ofParanthropus boisci. (RC 911 from Malema) known only from eastern Africa, represent the southernmost known distribution of these taxa. The biogeographic significance of these hominids from the Malawi-Rift lay in their association with the eastern African endemic animal group. Biogeographic variation in south-eastern Africa may be linked to habitat change occurring due to climate change, with maximum change occurring around 2.5 Ma.

In this chapter, the historical, systematic, and anatomical evidence for the diversity of the species within the australopith grade is reviewed. Given a strict evolutionary species definition, nominal taxonomic diversity and species‐lineage diversity do not necessarily map onto one another in the fossil record. Species‐lineages entail statements of ancestry and descent that depend on the consistency of phylogenetic and stratophenetic data. The requirements for identifying species‐lineages in the fossil record are severe, and in the early hominin record are rarely met, most often owing to small sample size, under‐ represented character data, nonrepresentation of rare or short‐lived taxa, poor chronological resolution, gaps in the time‐stratigraphic framework, or some combination of these factors. Because hypotheses concerning the ‘‘bushiness’’ of the hominin phylogenetic tree depend on the identification of lineages, not phenteically based ‘‘paleospecies,’’ confidence with respect to this issue is not justified for the majority of the hominin fossil record. There are two cases in which an approach to this question can be attempted. In one, the evidence is consistent with the evolution of Australopithecus anamensis into A. afarensis via anagenesis. The other, the evolution of A. boisei, most likely entailed a speciation event that gave rise to southern African clade (represented by A. robustus) subsequent to the appearance of A. aethiopicus. The late Pliocene time period in which the latter events transpired (ca. 2.8‐2.3 Ma) is one of substantial morphologicial diversity, high nominal taxonomic diversity, and high probability of synchronicity among known fossil samples. Although it is not possible to connect these australopith taxa (A. aethiopicus, A. africanus, A. garhi) to particular descendants due to defects in the data base, this time period probably documents a previously (and subsequently) unmatched degree of lineage proliferation compared to other parts of the human evolutionary record. The challenge to paleoanthropologists is to devote resources to improving this part of the fossil record and then to create testable phylogenetic and adaptive hypotheses to explain it.

Molecular studies suggest that the lineages leading to humans and chimpanzees diverged approximately 6.5–5.5 million years (Myr) ago, in the Late Miocene1,2,3. Hominid fossils from this interval, however, are fragmentary and of uncertain phylogenetic status, age, or both4,5,6. Here I report new hominid specimens from the Middle Awash area of Ethiopia that date to 5.2–5.8 Myr and are associated with a wooded palaeoenvironment7. These Late Miocene fossils are assigned to the hominid genus Ardipithecus and represent the earliest definitive evidence of the hominid clade. Derived dental characters are shared exclusively with all younger hominids. This indicates that the fossils probably represent a hominid taxon that postdated the divergence of lineages leading to modern chimpanzees and humans. However, the persistence of primitive dental and postcranial characters in these new fossils indicates that Ardipithecus was phylogenetically close to the common ancestor of chimpanzees and humans. These new findings raise additional questions about the claimed hominid status of Orrorin tugenensis8, recently described from Kenya and dated to ∼6 Myr9.

Remains of an early hominid have been recovered from four localities in the Lukeino Formation, Tugen Hills, Kenya, in sediments aged ca 6 Ma. 13 fossils are known, belonging to at least five individuals. The femora indicate that the Lukeino hominid was a biped when on the ground, whilst its humerus and manual phalanx show that it possessed some arboreal adaptations. The upper central incisor is large and robust, the upper canine is large for a hominid and retains a narrow and shallow anterior groove, the lower fourth premolar is ape-like, with offset roots and oblique crown, and the molars are relatively small, with thick enamel. A new genus and species is erected for the remains.

Over the past decade, discussions of the evolution of the earliest human ancestors have focused on the locomotion of the australopithecines. Recent discoveries in a broad range of disciplines have raised important questions about the influence of ecological factors in early human evolution. Here we trace the cranial and dental traits of the early australopithecines through time, to show that between 4.4 million and 2.3 million years ago, the dietary capabilities of the earliest hominids changed dramatically, leaving them well suited for life in a variety of habitats and able to cope with significant changes in resource availability associated with long-term and short-term climatic fluctuations.

This study uses the interpretation of paleosol features at Kanapoi, Kenya (4.2-3.4 Ma) to reconstruct the ecosystem occupied by Australopithecus anamensis. The paleosols at Kanapoi provide a unique and fortuitous opportunity, in that the bulk of the hominid specimens derive from paleosols, providing direct evidence of the environment that the Kanapoi hominids occupied. Seven named types of paleosols are recognized at Kanapoi, each representing a trace fossil of the local ecosystem during soil formation. The hominid-bearing Dite paleosols provide evidence that A. anamensis inhabited areas of semi-arid, seasonal climate regimes with mean annual precipitation ranging from about 350-600 mm. The in situ hominid collections from Dite paleosols show that A. anamensis at least occasionally occupied relatively open low tree-shrub savanna vegetation formed in well drained settings, and may have preferred these conditions over other poorly drained soils. The relatively open conditions of Dite paleosols existed within a spatially variable ecosystem, characterized by a mosaic of environments, ranging from forb-dominated edaphic grassland to gallery woodland, providing a larger view of the mixed ecosystem in which A. anamensis lived. Synthesis of paleoenvironmental indicators of A. anamensis at Kanapoi and Allia Bay, Kenya suggests that as early as 4 Ma hominids thrived in varied ecosystems.