Concept: Homo habilis
Homo naledi is a previously-unknown species of extinct hominin discovered within the Dinaledi Chamber of the Rising Star cave system, Cradle of Humankind, South Africa. This species is characterized by body mass and stature similar to small-bodied human populations but a small endocranial volume similar to australopiths. Cranial morphology of H. naledi is unique, but most similar to early Homo species including Homo erectus, Homo habilis or Homo rudolfensis. While primitive, the dentition is generally small and simple in occlusal morphology. H. naledi has humanlike manipulatory adaptations of the hand and wrist. It also exhibits a humanlike foot and lower limb. These humanlike aspects are contrasted in the postcrania with a more primitive or australopith-like trunk, shoulder, pelvis and proximal femur. Representing at least 15 individuals with most skeletal elements repeated multiple times, this is the largest assemblage of a single species of hominins yet discovered in Africa.
There is much debate on the dietary adaptations of the robust hominin lineages during the Pliocene-Pleistocene transition. It has been argued that the shift from C3 to C4 ecosystems in Africa was the main factor responsible for the robust dental and facial anatomical adaptations of Paranthropus taxa, which might be indicative of the consumption of fibrous, abrasive plant foods in open environments. However, occlusal dental microwear data fail to provide evidence of such dietary adaptations and are not consistent with isotopic evidence that supports greater C4 food intake for the robust clades than for the gracile australopithecines. We provide evidence from buccal dental microwear data that supports softer dietary habits than expected for P. aethiopicus and P. boisei based both on masticatory apomorphies and isotopic analyses. On one hand, striation densities on the buccal enamel surfaces of paranthropines teeth are low, resembling those of H. habilis and clearly differing from those observed on H. ergaster, which display higher scratch densities indicative of the consumption of a wide assortment of highly abrasive foodstuffs. Buccal dental microwear patterns are consistent with those previously described for occlusal enamel surfaces, suggesting that Paranthropus consumed much softer diets than previously presumed and thus calling into question a strict interpretation of isotopic evidence. On the other hand, the significantly high buccal scratch densities observed in the H. ergaster specimens are not consistent with a highly specialized, mostly carnivorous diet; instead, they support the consumption of a wide range of highly abrasive food items.
Besides Homo erectus (sensu lato), the eastern African fossil record of early Homo has been interpreted as representing either a single variable species, Homo habilis, or two species. In the latter case, however, there is no consensus over the respective groupings, and which of the two includes OH 7, the 1.8-million-year-old H. habilis holotype. This partial skull and hand from Olduvai Gorge remains pivotal to evaluating the early evolution of the Homo lineage, and by priority names one or other of the two taxa. However, the distorted preservation of the diagnostically important OH 7 mandible has hindered attempts to compare this specimen with other fossils. Here we present a virtual reconstruction of the OH 7 mandible, and compare it to other early Homo fossils. The reconstructed mandible is remarkably primitive, with a long and narrow dental arcade more similar to Australopithecus afarensis than to the derived parabolic arcades of Homo sapiens or H. erectus. We find that this shape variability is not consistent with a single species of early Homo. Importantly, the jaw morphology of OH 7 is incompatible with fossils assigned to Homo rudolfensis and with the A.L. 666-1 Homo maxilla. The latter is morphologically more derived than OH 7 but 500,000 years older, suggesting that the H. habilis lineage originated before 2.3 million years ago, thus marking deep-rooted species diversity in the genus Homo. We also reconstructed the parietal bones of OH 7 and estimated its endocranial volume. At between 729 and 824 ml it is larger than any previously published value, and emphasizes the near-complete overlap in brain size among species of early Homo. Our results clarify the H. habilis hypodigm, but raise questions about its phylogenetic relationships. Differences between species of early Homo appear to be characterized more by gnathic diversity than by differences in brain size, which was highly variable within all taxa.
Modern humans are characterized by specialized hand morphology that is associated with advanced manipulative skills. Thus, there is important debate in paleoanthropology about the possible cause-effect relationship of this modern human-like (MHL) hand anatomy, its associated grips and the invention and use of stone tools by early hominins. Here we describe and analyse Olduvai Hominin (OH) 86, a manual proximal phalanx from the recently discovered >1.84-million-year-old (Ma) Philip Tobias Korongo (PTK) site at Olduvai Gorge (Tanzania). OH 86 represents the earliest MHL hand bone in the fossil record, of a size and shape that differs not only from all australopiths, but also from the phalangeal bones of the penecontemporaneous and geographically proximate OH 7 partial hand skeleton (part of the Homo habilis holotype). The discovery of OH 86 suggests that a hominin with a more MHL postcranium co-existed with Paranthropus boisei and Homo habilis at Olduvai during Bed I times.
The estimation of body size among the earliest members of the genus Homo (2.4-1.5Myr [millions of years ago]) is central to interpretations of their biology. It is widely accepted that Homo ergaster possessed increased body size compared with Homo habilis and Homo rudolfensis, and that this may have been a factor involved with the dispersal of Homo out of Africa. The study of taxonomic differences in body size, however, is problematic. Postcranial remains are rarely associated with craniodental fossils, and taxonomic attributions frequently rest upon the size of skeletal elements. Previous body size estimates have been based upon well-preserved specimens with a more reliable species assessment. Since these samples are small (n < 5) and disparate in space and time, little is known about geographical and chronological variation in body size within early Homo. We investigate temporal and spatial variation in body size among fossils of early Homo using a 'taxon-free' approach, considering evidence for size variation from isolated and fragmentary postcranial remains (n = 39). To render the size of disparate fossil elements comparable, we derived new regression equations for common parameters of body size from a globally representative sample of hunter-gatherers and applied them to available postcranial measurements from the fossils. The results demonstrate chronological and spatial variation but no simple temporal or geographical trends for the evolution of body size among early Homo. Pronounced body size increases within Africa take place only after hominin populations were established at Dmanisi, suggesting that migrations into Eurasia were not contingent on larger body sizes. The primary evidence for these marked changes among early Homo is based upon material from Koobi Fora after 1.7Myr, indicating regional size variation. The significant body size differences between specimens from Koobi Fora and Olduvai support the cranial evidence for at least two co-existing morphotypes in the Early Pleistocene of eastern Africa.
The species Homo naledi was recently named from specimens recovered from the Dinaledi Chamber of the Rising Star cave system in South Africa. This large skeletal sample lacks associated faunal material and currently does not have a known chronological context. In this paper, we present comprehensive descriptions and metric comparisons of the recovered cranial and mandibular material. We describe 41 elements attributed to Dinaledi Hominin (DH1-DH5) individuals and paratype U.W. 101-377, and 32 additional cranial fragments. The H. naledi material was compared to Plio-Pleistocene fossil hominins using qualitative and quantitative analyses including over 100 linear measurements and ratios. We find that the Dinaledi cranial sample represents an anatomically homogeneous population that expands the range of morphological variation attributable to the genus Homo. Despite a relatively small cranial capacity that is within the range of australopiths and a few specimens of early Homo, H. naledi shares cranial characters with species across the genus Homo, including Homo habilis, Homo rudolfensis, Homo erectus, and Middle Pleistocene Homo. These include aspects of cranial form, facial morphology, and mandibular anatomy. However, the skull of H. naledi is readily distinguishable from existing species of Homo in both qualitative and quantitative assessments. Since H. naledi is currently undated, we discuss the evolutionary implications of its cranial morphology in a range of chronological frameworks. Finally, we designate a sixth Dinaledi Hominin (DH6) individual based on a juvenile mandible.
HWK EE is a little-known archaeological site from the top of Lower Bed II and the basal part of Middle Bed II, Olduvai Gorge, Tanzania. The site was originally excavated in the early 1970s by Mary Leakey, but the excavations and resulting lithic and fossil assemblages were never described. Here we report for the first time on the lithic and fossil assemblages that were recovered by Mary Leakey from the site. The lithic assemblage is one of the largest of any Oldowan site and is characterized by a core-and-flake technology with simple flaking techniques and minimal reduction of cores. Retouched flake frequencies and battered tools are higher than those reported for Olduvai Bed I and Lower Bed II assemblages, but flaking schemes are poorly organized. The fossil assemblage is well-preserved, taxonomically-rich, but dominated by bovids, and includes abundant feeding traces of both hominins and carnivores. Hominins are inferred to have broken the majority of limb bones at the site for access to marrow, while both carnivores and hominins likely had access to at least some flesh. HWK EE may represent one of the last Homo habilis sites at Olduvai Gorge, and is important to understanding the behavioral and cultural mechanisms that led to the emergence of the Acheulean and Homo erectus in the region.
The regular consumption of large mammal carcasses, as evidenced by butchery marks on fossils recovered from Early Stone Age archaeological sites, roughly coincides with the appearance of Homo habilis. However, the significance of this niche expansion cannot be appreciated without an understanding of hominin feeding behavior and their ecological interactions with mammalian carnivores. The Olduvai Geochronology and Archaeology Project (OGAP) has recovered a large and well-preserved fossil assemblage from the HWK EE site, which was deposited just prior to the first appearance of Acheulean technology at Olduvai Gorge and likely represents one of the last H. habilis sites at Olduvai. This taphonomic analysis of the larger mammal fossil assemblage excavated from HWK EE shows evidence of multiple occupations over a long period of time, suggesting the site offered resources that were attractive to hominins. There was a water source indicated by the presence of fish, crocodiles, and hippos, and there was possible tree cover in an otherwise open habitat. The site preserves several stratigraphic intervals with large fossil and artifact assemblages within two of these intervals. Feeding traces on bone surfaces suggest hominins at the site obtained substantial amounts of flesh and marrow, particularly from smaller size group 1-2 carcasses, and exploited a wide range of taxa, including megafauna. A strong carnivore signal suggests hominins scavenged much of their animal foods during the two main stratigraphic intervals. In the later interval, lower carnivore tooth mark and hammerstone percussion mark frequencies, in addition to high epiphyseal to shaft fragment ratios, suggest hominins and carnivores did not fully exploit bone marrow and grease, which may have been acquired from nutritionally-stressed animals that died during a dry period at Olduvai. The diversity of fauna that preserve evidence of butchery suggests that the HWK EE hominins were opportunistic in their acquisition of carcass foods.
Although the diminutive Homo floresiensis has been known for a decade, its phylogenetic status remains highly contentious. A broad range of potential explanations for the evolution of this species has been explored. One view is that H. floresiensis is derived from Asian Homo erectus that arrived on Flores and subsequently evolved a smaller body size, perhaps to survive the constrained resources they faced in a new island environment. Fossil remains of H. erectus, well known from Java, have not yet been discovered on Flores. The second hypothesis is that H. floresiensis is directly descended from an early Homo lineage with roots in Africa, such as Homo habilis; the third is that it is Homo sapiens with pathology. We use parsimony and Bayesian phylogenetic methods to test these hypotheses. Our phylogenetic data build upon those characters previously presented in support of these hypotheses by broadening the range of traits to include the crania, mandibles, dentition, and postcrania of Homo and Australopithecus. The new data and analyses support the hypothesis that H. floresiensis is an early Homo lineage: H. floresiensis is sister either to H. habilis alone or to a clade consisting of at least H. habilis, H. erectus, Homo ergaster, and H. sapiens. A close phylogenetic relationship between H. floresiensis and H. erectus or H. sapiens can be rejected; furthermore, most of the traits separating H. floresiensis from H. sapiens are not readily attributable to pathology (e.g., Down syndrome). The results suggest H. floresiensis is a long-surviving relict of an early (>1.75 Ma) hominin lineage and a hitherto unknown migration out of Africa, and not a recent derivative of either H. erectus or H. sapiens.
Among the ten fossil hominid deciduous teeth reported so far from the Pleistocene sediments of the Sangiran Dome of Java are two isolated lower second molars: specimens PCG.2 from the Kabuh Formation and FS-72 from the Pucangan Formation. While PCG.2 appears to be certainly attributable to Homo erectus, FS-72 is somewhat more problematic, even though it is commonly listed within the Indonesian H. erectus hypodigm. Largely because of its large size, it was originally attributed to Meganthropus paleojavanicus. Subsequent study highlighted a set of metric and nonmetric crown features also found in Australopith and African early Homo (notably H. habilis) homologues. An additional problem with the taxonomic assignment of isolated teeth from the Pleistocene of Java is the presence of Pongo in these same deposits.