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A partial skeleton from Hadar, Ethiopia (A.L. 438-1) attributed to Australopithecus afarensis is comprised of part of the mandible, a frontal bone fragment, a complete left ulna, two second metacarpals, one third metacarpal, plus parts of the clavicle, humerus, radius, and right ulna. It is one of only a few early hominin specimens to preserve both cranial and postcranial elements. It also includes the first complete ulna from a large A. afarensis individual, and the first associated metacarpal and forelimb remains. This specimen, dated to approximately 3Ma, is among the geologically youngest A. afarensis fossils and is also one of the largest individuals known. Its ulnar to mandibular proportions are similar to those of the geologically older and much smaller A.L. 288-1, suggesting that body size increased without disproportional enlargement of the mandible. Overall, however, analysis of this large specimen and of the diminutive A.L. 288-1 demonstrates that the functional morphology of the A. afarensis upper limb was similar at all body sizes; there is no evidence to support the hypothesis that more than one hominin species is present at Hadar. Morphologically, all apparent apomorphic traits of the elbow, forearm, wrist, and hand of A.L. 438-1 are shared uniquely with humans. Compared to humans, A.L. 438-1 does have a more curved ulna, although A.L. 288-1 does not, and it appears to have had slightly less well-developed manipulatory capabilities of its hands, although still more derived than in apes. We conclude that selection for effective arboreality in the upper limb of Australopithecus afarensis was weaker than in non-hominins, and that manipulative ability was of greater selective advantage than in extant great apes.

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The Hadar site in Ethiopia is a prolific source of hominid fossils attributed to the species Australopithecus afarensis, which spans the period 3.4-3.0 million years (myr) in the Sidi Hakoma, Denen Dora and lower Kada Hadar Members of the Hadar Formation. Since 1992 a major focus of field work conducted at Hadar has centered on sediments younger than 3.0 myr, comprising the bulk of the Kada Hadar Member. Witnessing the rise of the "robust" Australopithecus clade(s), the origin of Homo, and the first record of lithic artifacts, the period between 3.0 and 2.0 myr is strategically vital for paleoanthropology. However, in eastern Africa it is a particularly poorly sampled temporal interval. This paper provides a detailed comparative description of a hominid maxilla with partial dentition found at Hadar in 1994. The specimen, A.L. 666-1, derives from a lithic artifact-bearing horizon high in the Kada Hadar Member, 0.8 m below the BKT-3 tephra, dated by the 40Ar/39Ar method to 2.33 +/- 0.07 myr. Our preliminary investigation of the hominid specimen showed unambiguous affinities with early representatives of the Homo clade (Kimbel et al. [1996] J. Hum. Evol. 31:549-561). Further studies on maxillary and dental morphology lead us to attribute A.L. 666-1 to Homo aff. H. habilis. The new Hadar jaw is the first paleontological evidence for the projection of the H. habilis maxillofacial morphotype well back into the Pliocene. It may represent a male of this species, whose maxillary hypodigm consists chiefly of females. A subsidiary finding of our study is that of the three earliest recorded species of Homo (H. habilis, H. rudolfensis, H. erectus), it is H. habilis that exhibits facial morphology closest to that expected in their last common ancestor.

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The crescent of foramina of the cerebral surface of the sphenoid bone (superior orbital fissure, foramen rotundum, foramen ovale, foramen spinosum) differs morphologically in the African great apes and modern humans. New discoveries of Australopithecus afarensis at Hadar, Ethiopia, draw attention to the similarity of the crescent, particularly the "foramen" shape of the superior orbital fissure and its close proximity to the foramen rotundum, in this species, the African apes, and many other primates. Australopithecus africanus also shows this primitive pattern, whereas "robust" australopiths and humans share a configuration in which a true, laterally extended superior orbital fissure intervenes between the greater and lesser wings of the sphenoid and a broad bridge of bone separates the fissure from the foramen rotundum. This shared morphology may be added to the list of putative "robust" australopith-Homo synapomorphies.

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The Hadar Formation in Ethiopia is a prolific source of Pliocene Hominidae attributed to the species Australopithecus afarensis. Since 1990, three seasons of field work have contributed 53 new specimens to the hominid inventory from Hadar, including the first fairly complete adult skull. Ranging from 3.0 to 3.4 million years in age (Fig. 1), the new specimens bear on key debates in hominid palaeontology, including the taxonomic implications of sample variation and the reconstruction of locomotor behaviour. They confirm the taxonomic unity of A. afarensis and constitute the largest body of evidence for about 0.9 million years of stasis in the earliest known hominid species.

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The extensive overlap of the temporal squama on the parietal in KNM-WT 17000 is restricted to a narrow segment of the arc of the squamosal suture. It appears as a long, narrow, posterosuperior extension onto the calvarial wall. The long axis of this extended strip is aligned with what appears to be the most hypertrophied section of the temporalis muscle fan. The substantial differences between the squamosal suture and associated masticatory elements of KNM-WT 17000 and those of Australopithecus boisei provide an opportunity to evaluate the various forces molding the anatomy of the squamosal suture of A. boisei. It is suggested that the extreme flare of the zygomatic arches in the latter accounts for these differences.

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The recovery of a fossil hominid skeleton with a complete hyoid bone from Mousterian deposits in Kebara Cave, Israel, provides new evidence pertaining to the evolution of speech. Previous studies of speech in the Middle Palaeolithic (most notably those on Neandertals) have focused on the basicranium as an indicator of speech capabilities. This work critiques the use of the basicranium and instead presents the anatomical relations of the hyoid and adjacent structures in living humans as a basis for understanding the form of the vocal tract. The size and morphology of the hyoid from Kebara and its relations to other anatomical components are almost identical to those in modern humans, suggesting that Middle Palaeolithic populations were anatomically capable of fully modern speech.

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Two pelvises from a similar archaeological context have been discovered in recent years in two different caves in Israel. The pelvis from the Qafzeh cave (Qafzeh 9) was dated by means of thermoluminescence at approximately 95 kyr BP. All available measurement values, the most significant being those of the diagnostic obturator region, fall within those of the modern range. The other pelvis emanates from the Kebara cave and differs fundamentally from modern pelvises and from the Qafzeh specimen, although the Kebara pelvis is 30,000 years younger than the latter. As in other remains of Neandertal pelvises, the superior pubic ramus of the Kebara hominid is extremely long and slender and exhibits a cross section unlike that of modern humans. The absolute height of the obturator region is very small. It is these measurements and proportions that set the Kebara pelvis apart from both modern pelvises and the specimen from Qafzeh. The morphological differences and the chronological relationship between the two fossil pelvises support the concept of two distinct evolutionary lineages for these hominids.

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The origin of human language, and in particular the question of whether or not Neanderthal man was capable of language/speech, is of major interest to anthropologists but remains an area of great controversy. Despite palaeoneurological evidence to the contrary, many researchers hold to the view that Neanderthals were incapable of language/speech, basing their arguments largely on studies of laryngeal/basicranial morphology. Studies, however, have been hampered by the absence of unambiguous fossil evidence. We now report the discovery of a well-preserved human hyoid bone from Middle Palaeolithic layers of Kebara Cave, Mount Carmel, Israel, dating from about 60,000 years BP. The bone is almost identical in size and shape to the hyoid of present-day populations, suggesting that there has been little or no change in the visceral skeleton (including the hyoid, middle ear ossicles, and inferentially the larynx) during the past 60,000 years of human evolution. We conclude that the morphological basis for human speech capability appears to have been fully developed during the Middle Palaeolithic.

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The renewed excavations at the Kebara Cave revealed a Neanderthal skeleton dated at about 50-55,000 years B.P. The pelvis of this individual is the most intact Neanderthal pelvis yet discovered, presenting for the first time a complete inlet. Although the superior pubic ramus is extremely long, as typically seen in the Neanderthals, the size of the pelvic inlet is comparable to that of modern Homo sapiens. The length of the superior pubic ramus is found to stem from a more externally rotated innominate bone and not, as generally assumed, from the larger pelvic inlet. It is suggested that the uniqueness of the Neanderthal pelvis may be attributable to locomotion and posture-related biomechanics rather than to obstetric requirements.

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The beginning of specialization characterizing the robust australopithecines is manifested in almost every aspect of the masticatory system of Australopithecus africanus. Of particular significance is the presence of two massive bony columns on both sides of the nasal aperture that support the anterior portion of the palate. These columns--the anterior pillars--are viewed as a structural response to the greater occlusal load stemming from the beginning stages of molarization of the premolars and exerted on the more anterior part of the dental arcade. In A. africanus the molarization process is, indeed, just in its initial phase, but the still considerable protrusion of the palate relative to the more peripheral facial frame increases the need for pillars. The anterior pillars and the advancement of the inferior part of the infraorbital plate (the origin of the masseter) play a major role in molding the facial topography of A. africanus. The absence of the pillars and the common position of the masseteric origin lead us to define the face of A. afarensis as the most primitive of the australopithecines and allow us to discriminate between its facial morphology and that of A. africanus. The presence of anterior pillars in the face of the latter places it clearly in the robust australopithecine clade.

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Asterionic sutural patterns in Plio-Pleistocene hominid crania have never been examined in detail. We present an analysis of this anatomical region in Australopithecus and Homo and relate different sutural patterns to functional changes in the masticatory apparatus. The great apes and A. afarensis share the common adult higher primate sutural pattern referred to as the "asterionic notch," which develops in response to the hypertrophy of posterior temporalis muscle fibers and the consequent formation of compound temporal/nuchal crests. This sutural configuration also appears to be present on the early Homo cranium KNM-ER 1805. In contrast, adult male A. boisei crania exhibit a unique pattern where the temporal squama overlaps the parietal which, in turn, overlaps the par mastoidea and the upper scale of the occipital bone. We relate this arrangement to the need to reinforce the rear of a thin-walled braincase against the net tensile forces exerted by the temporalis and nuchal muscles. The common juvenile hominoid edge-to-edge asterionic articulation is maintained in adult A. africanus, A. robustus, female A. boisei, and most Homo crania. We discuss the latter pattern in regard to anterior temporalis hypertrophy in A. africanus, A. robustus, and A. boisei and to craniofacial paedomorphosis in Homo.

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The nature of human evolution has been viewed recently as a specific example of a more general model of evolution termed "punctuated equilibrium". The characteristics of this model are long periods of little or not evolutionary change (stasis) interspersed with periods of rapid (punctuated) morphological change. Careful analysis of the hominid fossil record over the past 4.0 million years, however, suggests no well documented examples of either stasis or punctuation. The evidence for the evolution of the hominid lineage is most reasonably interpreted by a model of more gradual change with periods of varying rates of evolution.

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A brief description of the exposed middle ear from two Swartkrans specimens demonstrates the feasibility of examining the inner auditory structures of existing fossil hominid material. These Swartkrans fossils provide the first clear view of the early hominid middle ear, and their structures appear to be morphologically distinct. The bony anatomy of the external ear of Australopithecus robustus is found to be unique among the South African hominids.

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A juvenile Australopithecus boisei specimen from the Omo basin, southern Ethiopia, is found to exhibit and extraordinarily large overlap of the temporal squama on the parietal, a phenomenon shared with at least two adult specimens of A. boisei. An attempt is made to interpret the overlap as a structural (bony/ligamentous) adaptation necessitated by the unique combination of certain components of the masticatory system of A. boisei. These are: (1) the massiveness and strength of the temporalis muscle, (2) its relatively anterior location, and (3) the lateral position of the masseter muscle due to the flaring of the zygomatic arches. The effect of the temporalis muscle is to create excessive pressure on the portion of the squamosal suture along the parietal, while the lateral placement of the masseter and the resultant increase of pressure on the temporal squama via the zygomatic arch tend to "loosen" the contact between the temporal and parietal bones.

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