An ancient cranium from Dmanisi: Evidence for interpersonal violence, disease, and possible predation by carnivores on Early Pleistocene Homo.

Five well-preserved individuals from Dmanisi represent a paleodeme attributed to early Homo. Here we provide a case study of the D2280 adult cranium, which presents four oval-shaped lesions on the frontal, left parietal, and occipital bones. Several conditions are considered as possibly contributing to this pathology, including trauma, cysts, metastatic cancer, and infectious disease. One large but shallow depression on the left parietal bone has slightly elevated boundaries. Imaging reveals inner and outer tables that are reciprocally concave, so that the diploë is diminished or completely absent. This lesion is very likely a result of traumatic injury. Two additional depressions on the left side frontal and occipital bones may also be attributed to blunt force trauma. Such injuries stem from a variety of causes, but interpersonal violence may well be implicated. Based on the location and structure of a fourth lesion on the right-side frontal bone, we advance a possible diagnosis of treponemal disease. Lesions on the cranium and specifically on the frontal bone are common in treponemal disease. The condition develops as a periostitis, which eventually results in the destruction of the osteoperiosteal border of the cranial outer table and rarely involves the inner table. Additional perforations on the occipital bone are interpreted as perimortem damage resulting from predation by carnivores or scavenging by birds. If our reading of this cranial evidence can be confirmed, then D2280 documents one of the earliest instances of blunt force trauma in the Homo lineage. Dmanisi may also reveal the presence of treponemal disease in a population dated ca. 1.77 Ma ago. These findings bear on the social behavior of ancient humans and also the impact of infectious diseases on their survival.

Variation among the Dmanisi hominins: Multiple taxa or one species?

OBJECTIVES: There is continuing controversy over the number of taxa documented by the Dmanisi hominins. Variation may reflect age and sex differences within a single population. Alternatively, two (or more) distinct species may be present. Our null hypothesis states that just one population is represented at the site. MATERIALS AND METHODS: We assess the likely sources of variation in endocranial capacity, craniofacial and mandibular morphology, and the expression of characters related to aging and sex dimorphism. We use the coefficient of variation and a modified version of Levene's test for equal variances to compare trait variation at Dmanisi with that in fossil hominins and modern Homo sapiens from Africa. RESULTS: Skull 5 presents a low, massive vault, and a muzzle-like lower face. Other individuals have larger brains and more globular vaults. Despite such variation, the five crania share numerous features. All of the mandibles possess marginal tubercles, mandibular tori, and a distinctive patterning of mental foramina. Relative variation at Dmanisi is comparable to that in selected reference groups. Further growth anticipated in Skull 3, age-related remodeling affecting the D2600 mandible, pathology, and sex dimorphism can account for much of the interindividual variation observed. The preponderance of evidence supports our null hypothesis. DISCUSSION: Sources of the variation within ancient Homo assemblages remain poorly understood. Skull 5 is a very robust male, with a brain smaller than that of both a juvenile (Skull 3) and a probable female (Skull 2). Skull 1 has the largest brain, but cranial superstructures do not clearly mark this individual as male or female. It is likely that the Dmanisi hominins represent a single paleospecies of Homo displaying a pattern of sex dimorphism not seen in living hominids.

Did Our Species Evolve in Subdivided Populations across Africa, and Why Does It Matter?

We challenge the view that our species, Homo sapiens, evolved within a single population and/or region of Africa. The chronology and physical diversity of Pleistocene human fossils suggest that morphologically varied populations pertaining to the H. sapiens clade lived throughout Africa. Similarly, the African archaeological record demonstrates the polycentric origin and persistence of regionally distinct Pleistocene material culture in a variety of paleoecological settings. Genetic studies also indicate that present-day population structure within Africa extends to deep times, paralleling a paleoenvironmental record of shifting and fractured habitable zones. We argue that these fields support an emerging view of a highly structured African prehistory that should be considered in human evolutionary inferences, prompting new interpretations, questions, and interdisciplinary research directions.

Skull 5 from Dmanisi: Descriptive anatomy, comparative studies, and evolutionary significance.

A fifth hominin skull (cranium D4500 and mandible D2600) from Dmanisi is massively constructed, with a large face and a very small brain. Traits documented for the first time in a basal member of the Homo clade include the uniquely low ratio of endocranial volume to basicranial width, reduced vertex height, angular vault profile, smooth nasal sill coupled with a long and sloping maxillary clivus, elongated palate, and tall mandibular corpus. The convex clivus and receding symphysis of skull 5 produce a muzzle-like form similar to that of Australopithecus afarensis. While the Dmanisi cranium is very robust, differing from OH 13, OH 24, and KNM-ER 1813, it resembles Homo habilis specimens in the "squared off" outline of its maxilla in facial view, maxillary sulcus, rounded and receding zygomatic arch, and flexed zygomaticoalveolar pillar. These characters distinguish early Homo from species of Australopithecus and Paranthropus. Skull 5 is unlike Homo rudolfensis cranium KNM-ER 1470. Although it appears generally primitive, skull 5 possesses a bar-like supraorbital torus, elongated temporal squama, occipital transverse torus, and petrotympanic traits considered to be derived for Homo erectus. As a group, the Dmanisi crania and mandibles display substantial anatomical and metric variation. A key question is whether the fossils document age-related growth and sex dimorphism within a single population, or whether two (or more) distinct taxa may be present at the site. We use the coefficient of variation to compare Dmanisi with Paranthropus boisei, H. erectus, and recent Homo sapiens, finding few signals that the Dmanisi sample is excessively variable in comparison to these reference taxa. Using cranial measurements and principal components analysis, we explore the proposal that the Dmanisi skulls can be grouped within a regionally diverse hypodigm for H. erectus. Our results provide only weak support for this hypothesis. Finally, we consider all available morphological and paleobiological evidence in an attempt to clarify the phyletic relationship of Dmanisi to Homo species evolving >2.0 to 1.0 Ma.

Response to comment on "A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo".

Schwartz et al. hold that variation among the Dmanisi skulls reflects taxic diversity. The morphological observations to support their hypothesis, however, are partly incorrect, and not calibrated against intraspecific variation in living taxa. After proper adjustment, Schwartz et al.'s data are fully compatible with the hypothesis of a single paleodeme of early Homo at Dmanisi.

A complete skull from Dmanisi, Georgia, and the evolutionary biology of early Homo.

The site of Dmanisi, Georgia, has yielded an impressive sample of hominid cranial and postcranial remains, documenting the presence of Homo outside Africa around 1.8 million years ago. Here we report on a new cranium from Dmanisi (D4500) that, together with its mandible (D2600), represents the world's first completely preserved adult hominid skull from the early Pleistocene. D4500/D2600 combines a small braincase (546 cubic centimeters) with a large prognathic face and exhibits close morphological affinities with the earliest known Homo fossils from Africa. The Dmanisi sample, which now comprises five crania, provides direct evidence for wide morphological variation within and among early Homo paleodemes. This implies the existence of a single evolving lineage of early Homo, with phylogeographic continuity across continents.

Homo erectus and Middle Pleistocene hominins: brain size, skull form, and species recognition.

Hominins that differ from Homo erectus, the Neanderthals, and recent humans are known from Middle Pleistocene localities across the Old World. The taxonomic status of these populations has been clouded by controversy. Perhaps the most critical problem has been an incomplete understanding of variation in skull form. Here, both H. erectus and later mid-Pleistocene hominins are the focus of an investigation aimed at clarifying the relationships among brain volume, basicranial dimensions, neurocranial shape, and certain facial characters. Brain size in H. erectus averages about 950 cm(3), while in a series of Middle Pleistocene crania from Africa and Europe, volume is about 1230 cm(3). If encephalization is the primary mechanism operating in the mid-Pleistocene, then diverse aspects of cranial form cannot all be treated as independent variables. Correlation is utilized to examine the associations among measurements for more than 30 H. erectus crania that are reasonably well preserved. A similar approach is used with the Middle Pleistocene sample. Patterns of covariation are compared in order to assess integration. Next, factor analysis is applied to the H. erectus specimens in an attempt to identify modules, tightly integrated traits that can evolve independently. Studies of the variation within H. erectus are followed by direct comparisons with the Middle Pleistocene population. Discriminant functions facilitate the description of intergroup differences. Traits that vary independently from brain volume include anterior frontal broadening, lateral expansion of the parietal vault, elevation of the lambda-inion chord, and rounding of the sagittal contour of the occipital. This finding helps to resolve the problem of species recognition. Neurocranial proportions as well as characters from the cranial base and face can be incorporated into a differential diagnosis for the mid-Pleistocene sample. Evidence presented here supports arguments for speciation in the Middle Pleistocene.

Radiometric dating of the type-site for Homo heidelbergensis at Mauer, Germany.

The Mauer mandible, holotype of Homo heidelbergensis, was found in 1907 in fluvial sands deposited by the Neckar River 10 km southeast of Heidelberg, Germany. The fossil is an important key to understanding early human occupation of Europe north of the Alps. Given the associated mammal fauna and the geological context, the find layer has been placed in the early Middle Pleistocene, but confirmatory chronometric evidence has hitherto been missing. Here we show that two independent techniques, the combined electron spin resonance/U-series method used with mammal teeth and infrared radiofluorescence applied to sand grains, date the type-site of Homo heidelbergensis at Mauer to 609 ± 40 ka. This result demonstrates that the mandible is the oldest hominin fossil reported to date from central and northern Europe and raises questions concerning the phyletic relationship of Homo heidelbergensis to more ancient populations documented from southern Europe and in Africa. We address the paleoanthropological significance of the Mauer jaw in light of this dating evidence.

Locomotor anatomy and biomechanics of the Dmanisi hominins.

The Dmanisi hominins inhabited a northern temperate habitat in the southern Caucasus, approximately 1.8 million years ago. This is the oldest population of hominins known outside of Africa. Understanding the set of anatomical and behavioral traits that equipped this population to exploit their seasonal habitat successfully may shed light on the selection pressures shaping early members of the genus Homo and the ecological strategies that permitted the expansion of their range outside of the African subtropics. The abundant stone tools at the site, as well as taphonomic evidence for butchery, suggest that the Dmanisi hominins were active hunters or scavengers. In this study, we examine the locomotor mechanics of the Dmanisi hind limb to test the hypothesis that the inclusion of meat in the diet is associated with an increase in walking and running economy and endurance. Using comparative data from modern humans, chimpanzees, and gorillas, as well as other fossil hominins, we show that the Dmanisi hind limb was functionally similar to modern humans, with a longitudinal plantar arch, increased limb length, and human-like ankle morphology. Other aspects of the foot, specifically metatarsal morphology and tibial torsion, are less derived and similar to earlier hominins. These results are consistent with hypotheses linking hunting and scavenging to improved walking and running performance in early Homo. Primitive retentions in the Dmanisi foot suggest that locomotor evolution continued through the early Pleistocene.

Out of Africa: modern human origins special feature: middle and later Pleistocene hominins in Africa and Southwest Asia.

Approximately 700,000 years ago, Homo erectus in Africa was giving way to populations with larger brains accompanied by structural adjustments to the vault, cranial base, and face. Such early Middle Pleistocene hominins were not anatomically modern. Their skulls display strong supraorbital tori above projecting faces, flattened frontals, and less parietal expansion than is the case for Homo sapiens. Postcranial remains seem also to have archaic features. Subsequently, some groups evolved advanced skeletal morphology, and by ca. 200,000 years ago, individuals more similar to recent humans are present in the African record. These fossils are associated with Middle Stone Age lithic assemblages and, in some cases, Acheulean tools. Crania from Herto in Ethiopia carry defleshing cutmarks and superficial scoring that may be indicative of mortuary practices. Despite these signs of behavioral innovation, neither the Herto hominins, nor others from Late Pleistocene sites such as Klasies River in southern Africa and Skhul/Qafzeh in Israel, can be matched in living populations. Skulls are quite robust, and it is only after approximately 35,000 years ago that people with more gracile, fully modern morphology make their appearance. Not surprisingly, many questions concerning this evolutionary history have been raised. Attention has centered on systematics of the mid-Pleistocene hominins, their paleobiology, and the timing of dispersals that spread H. sapiens out of Africa and across the Old World. In this report, I discuss structural changes characterizing the skulls from different time periods, possible regional differences in morphology, and the bearing of this evidence on recognizing distinct species.

Postcranial evidence from early Homo from Dmanisi, Georgia.

The Plio-Pleistocene site of Dmanisi, Georgia, has yielded a rich fossil and archaeological record documenting an early presence of the genus Homo outside Africa. Although the craniomandibular morphology of early Homo is well known as a result of finds from Dmanisi and African localities, data about its postcranial morphology are still relatively scarce. Here we describe newly excavated postcranial material from Dmanisi comprising a partial skeleton of an adolescent individual, associated with skull D2700/D2735, and the remains from three adult individuals. This material shows that the postcranial anatomy of the Dmanisi hominins has a surprising mosaic of primitive and derived features. The primitive features include a small body size, a low encephalization quotient and absence of humeral torsion; the derived features include modern-human-like body proportions and lower limb morphology indicative of the capability for long-distance travel. Thus, the earliest known hominins to have lived outside of Africa in the temperate zones of Eurasia did not yet display the full set of derived skeletal features.

A fourth hominin skull from Dmanisi, Georgia.

Newly discovered Homo remains, stone artifacts, and animal fossils from Dmanisi, Republic of Georgia, provide a basis for better understanding patterns of hominin evolution and behavior in Eurasia ca. 1.77 million years ago. Here we describe a fourth skull that is nearly complete, lacking all but one of its teeth at the time of death. Both the maxillae and the mandible exhibit extensive bone loss due to resorption. This individual is similar to others from the site but supplies information about variation in brain size and craniofacial anatomy within the Dmanisi paleodeme. Although this assemblage presents numerous primitive characters, the Dmanisi skulls are best accommodated within the species H. erectus. On anatomical grounds, it is argued that the relatively small-brained and lightly built Dmanisi hominins may be ancestral to African and Far Eastern branches of H. erectus showing more derived morphology.

Human foot bones from Klasies River main site, South Africa.

The caves at Klasies River contain abundant archaeological evidence relating to human evolution in the late Pleistocene of southern Africa. Along with Middle Stone Age artifacts, animal bones, and other food waste, there are hominin cranial fragments, mandibles with teeth, and a few postcranial remains. Three foot bones can now be added to this inventory. An adult first metatarsal is similar in size and discrete anatomical features to those from Holocene burials in the Cape Province. A complete and well-preserved second metatarsal is especially long and heavy at midshaft in comparison to all Holocene and more recent South African homologues. A large fifth metatarsal is highly distinctive in its morphology. In overall size, these pedal elements resemble specimens from late Pleistocene sites in western Asia, but there are some differences in proportions. The fossils support earlier suggestions concerning a relatively high level of sexual dimorphism in the African Middle Stone Age population. Squatting facets on the two lateral metatarsals appear to indicate a high frequency of kneeling among members of this group. The new postcranial material also underlines the fact that the morphology of particular skeletal elements of some of the 100,000-year-old Klasies River individuals falls outside the range of modern variation.

Anatomical descriptions, comparative studies and evolutionary significance of the hominin skulls from Dmanisi, Republic of Georgia.

Evidence for ancient hominin occupation in Eurasia comes from Dmanisi in the Georgian Caucasus. Stratigraphic and sedimentological arguments, geochemical observations, paleomagnetic sampling and radiometric dates all point to the conclusion that bones and artifacts were deposited at this site during a brief interval following the close of the Olduvai Subchron (1.77 million years ago). In this report we present further descriptive and comparative studies of the D2280 braincase, the D2282 partial cranium, now linked with the D211 mandible, and the skull D2700/D2735. The crania have capacities ranging from 600 cm3 to 775 cm3. Supraorbital tori and other vault superstructures are only moderately developed. The braincase is expanded laterally in the mastoid region, but the occiput is rounded. The pattern of sagittal keeling is distinctive. D2700 displays a prominent midfacial profile and has a very short nasoalveolar clivus. Also, the M3 crowns are reduced in size. Although there is variation probably related to growth status and sex dimorphism, it is appropriate to group the Dmanisi hominins together. With the possible exception of the large D2600 mandible, the individuals are sampled from one paleodeme. This population resembles Homo habilis in brain volume and some aspects of craniofacial morphology, but many of these features can be interpreted as symplesiomorphies. Other discrete characters and measurements suggest that the Dmanisi skulls are best placed with H. erectus. There are numerous similarities to individuals from the Turkana Basin in Kenya, but a few features link Dmanisi to Sangiran in Java. Some traits expressed in the Dmanisi assemblage appear to be unique. Reconstructing the evolutionary relationships of these ancient populations of Africa and Eurasia is difficult, as the record is quite patchy, and determination of character polarities is not straightforward. Nevertheless, the evidence from anatomical analysis and measurements supports the hypothesis that Dmanisi is close to the stem from which H. erectus evolved.

Anthropology: the earliest toothless hominin skull.

The site of Dmanisi in the Eurasian republic of Georgia has yielded striking hominin, faunal and archaeological material as evidence for the presence of early Homo outside Africa 1.77 million years ago, documenting an important episode in human evolution. Here we describe a beautifully preserved skull and jawbone from a Dmanisi hominin of this period who had lost all but one tooth several years before death. This specimen not only represents the earliest case of severe masticatory impairment in the hominin fossil record to be discovered so far, but also raises questions about alternative subsistence strategies in early Homo.

Brain size and encephalization in early to Mid-Pleistocene Homo.

Important changes in the brain have occurred during the course of human evolution. Both absolute and relative size increases can be documented for species of Homo, culminating in the appearance of modern humans. One species that is particularly well-represented by fossil crania is Homo erectus. The mean capacity for 30 individuals is 973 cm(3). Within this group there is substantial variation, but brain size increases slightly in specimens from later time periods. Other Middle Pleistocene crania differ from those of Homo erectus. Characters of the facial skeleton, vault, and cranial base suggest that fossils from sites such as Arago Cave in France, the Sima de los Huesos in Spain, Bodo in Ethiopia, Broken Hill in Zambia, and perhaps Dali in China belong to the taxon Homo heidelbergensis. Ten of these mid-Quaternary hominins have brains averaging 1,206 cm(3) in volume, and many fall beyond the limits of size predicted for Homo erectus of equivalent age. When orbit height is used to construct an index of relative brain size, it is apparent that the (significant) increase in volume documented for the Middle Pleistocene individuals is not simply a consequence of larger body mass. Encephalization quotient values confirm this finding. These changes in absolute and relative brain size can be taken as further corroborative evidence for a speciation event, in which Homo erectus produced a daughter lineage. It is probable that Homo heidelbergensis originated in Africa or western Eurasia and then ranged widely across the Old World. Archaeological traces indicate that these populations differed in their technology and behavior from earlier hominins.

A new skull of early Homo from Dmanisi, Georgia.

Another hominid skull has been recovered at Dmanisi (Republic of Georgia) from the same strata in which hominid remains have been reported previously. The Dmanisi site dated to approximately 1.75 million years ago has now produced craniofacial portions of several hominid individuals, along with many well-preserved animal fossils and quantities of stone artifacts. Although there are certain anatomical differences among the Dmanisi specimens, the hominids do not clearly represent more than one taxon. We assign the new skull provisionally to Homo erectus (=ergaster). The Dmanisi specimens are the most primitive and small-brained fossils to be grouped with this species or any taxon linked unequivocally with genus Homo and also the ones most similar to the presumed habilis-like stem. We suggest that the ancestors of the Dmanisi population dispersed from Africa before the emergence of humans identified broadly with the H. erectus grade.