Search

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].

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).

International audience; A fossil hominin cranium was discovered in mid-Pliocene deltaic strata in the Godaya Valley of the northwestern Woranso-Mille study area in Ethiopia. Here we show that analyses of chemically correlated volcanic layers and the palaeomagnetic stratigraphy, combined with Bayesian modelling of dated tuffs, yield an age range of 3.804 ± 0.013 to 3.777 ± 0.014 million years old (mean ± 1σ) for the deltaic strata and the fossils that they contain. We also document deposits of a perennial lake beneath the deltaic sequence. Mammalian fossils associated with the cranium represent taxa that were widespread at the time and data from botanical remains indicate that the vegetation in the lake and delta catchment was predominantly dry shrubland with varying proportions of grassland, wetland and riparian forest. In addition, we report high rates of sediment accumulation and depositional features that are typical of a steep topographic relief and differ from younger Woranso-Mille fossil localities, reflecting the influence of active rift processes on the palaeolandscape. Q1 Until recently, field work in the Woranso-Mille area (Afar, Ethiopia) has concentrated on exposures along the Mille River that include more than 120 m of fossiliferous sedimentary and volcanic strata that are between 3.8 and 3.2 million years (Myr) old. East of Korsi Dora (Fig. 1a), sedimentary sequences that are younger than the extra-regional approximately 3.57-Myr-old Kilaytoli tuff 1 have yielded specimens of at least two hominin species, Australopithecus afarensis and Australopithecus deyiremeda 2-5

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.

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.

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.

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.

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.

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.

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.

International audience; Until recently, our understanding of the evolution of human growth and development derived from studies of fossil juveniles that employed extant populations for both age determination and comparison. This circular approach has led to considerable debate about the humanlike and ape-like affinities of fossil hominins. Teeth are invaluable for understanding maturation as age at death can be directly assessed from dental microstructure, and dental development has been shown to correlate with life history across primates broadly. We employ non-destructive synchrotron imaging to characterize incremental development, molar emergence, and age at death in more than 20 Australopithecus anamensis, Australopithecus africanus, Paranthropus robustus and South African early Homo juveniles. Long-period line periodicities range from at least 6-12 days (possibly 5-13 days), and do not support the hypothesis that australopiths have lower mean values than extant or fossil Homo. Crown formation times of australopith and early Homo postcanine teeth fall below or at the low end of extant human values; Paranthropus robustus dentitions have the shortest formation times. Pliocene and early Pleistocene hominins show remarkable variation, and previous reports of age at death that employ a narrow range of estimated long-period line periodicities, cuspal enamel thicknesses, or initiation ages are likely to be in error. New chronological ages for SK 62 and StW 151 are several months younger than previous histological estimates, while Sts 24 is more than one year older. Extant human standards overestimate age at death in hominins predating Homo sapiens, and should not be applied to other fossil taxa. We urge caution when inferring life history as aspects of dental development in Pliocene and early Pleistocene fossils are distinct from modern humans and African apes, and recent work has challenged the predictive power of primate-wide associations between hominoid first molar emergence and certain life history variables

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.

Understanding changes in ontogenetic development is central to the study of human evolution. With the exception of Neanderthals, the growth patterns of fossil hominins have not been studied comprehensively because the fossil record currently lacks specimens that document both cranial and postcranial development at young ontogenetic stages. Here we describe a well-preserved 3.3-million-year-old juvenile partial skeleton of Australopithecus afarensis discovered in the Dikika research area of Ethiopia. The skull of the approximately three-year-old presumed female shows that most features diagnostic of the species are evident even at this early stage of development. The find includes many previously unknown skeletal elements from the Pliocene hominin record, including a hyoid bone that has a typical African ape morphology. The foot and other evidence from the lower limb provide clear evidence for bipedal locomotion, but the gorilla-like scapula and long and curved manual phalanges raise new questions about the importance of arboreal behaviour in the A. afarensis locomotor repertoire.

International audience; The origin of Australopithecus, the genus widely interpreted as ancestral to Homo, is a central problem in human evolutionary studies. Australopithecus species differ markedly from extant African apes and candidate ancestral hominids such as Ardipithecus, Orrorin and Sahelanthropus. The earliest described Australopithecus species is Au. anamensis, the probable chronospecies ancestor of Au. afarensis. Here we describe newly discovered fossils from the Middle Awash study area that extend the known Au. anamensis range into northeastern Ethiopia. The new fossils are from chronometrically controlled stratigraphic sequences and date to about 4.1–4.2 million years ago. They include diagnostic craniodental remains, the largest hominid canine yet recovered, and the earliest Australopithecus femur. These new fossils are sampled from a woodland context. Temporal and anatomical intermediacy between Ar. ramidus and Au. afarensis suggest a relatively rapid shift from Ardipithecus to Australopithecus in this region of Africa, involving either replacement or accelerated phyletic evolution.