Human evolution

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or ‘early African Homo erectus’, which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of ‘archaic Homo’ in Europe is dated at between 600–700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from ‘archiac’ to ‘modern’ Homo may have taken place in Africa.     

L’architecture de l’épaule au sein du genre Homo : nouvelles interprétations

The morphology of human clavicles can be estimated by projecting them on two perpendicular planes in order to assess the shapes of their cranial and dorsal primary curvatures. In cranial view no differences in curvature appear within the genus Homo, which means the different species had similar arms elevation capacity, especially in protraction. On the contrary, in dorsal view two clavicles morphologies could be defined. The first one is characterized by two curvatures in dorsal view and is possessed by all Homo species, from Homo habilis to Neanderthal, including Homo ergaster, but not modern human, Upper Paleolithic and anatomically modern human remains, who possess clavicles of the second type, characterized by either one curvature, or two slightly pronounced ones in dorsal view. Clavicles displaying two pronounced curvatures in dorsal view are associated with scapula sitting high on the thorax in regard to modern human. However, shoulder with high scapula on the thorax displays two different kinds of architectures: (i) shoulder with short clavicles associated to scapulas sitting more laterally than those of modern human. This group includes earlier Homo like Homo habilis and Homo ergaster and (ii) shoulder with long clavicles associated to scapulas sitting more dorsally on the thorax, like those of modern human. This group includes Homoantecessor and Neanderthals. In other words, within the genus Homo, three shoulders would have existed. Evolution of the shoulder complex is far more complex than previously thought and the arrival of modern bipedalism was not associated to modern shoulder.     

Origin of the Genus <Emphasis Type="Italic">Homo</Emphasis>

The origin of the genus Homo in Africa signals the beginning of the shift from increasingly bipedal apes to primitive, large-brained, stone tool-making, meat-eaters that traveled far and wide. This early part of the human genus is represented by three species: Homo habilis, Homo rudolfensis, and Homo erectus. H. habilis is known for retaining primitive features that link it to australopiths and for being the first stone tool makers. Little is known about H. rudolfensis except that it had a relatively large brain and large teeth compared to H. habilis and that it overlapped in time and space with other early Homo. Our understanding of the paleobiology and evolution of the larger-brained H. erectus is enhanced due to its rich fossil record. H. erectus was the first obligate, fully committed biped, and with a body adapted for modern striding locomotion, it was also the first in the human lineage to disperse outside of Africa. The early members of the genus Homo are the first to tip the scale from the more apish side of our evolutionary history toward the more human one.     

Emergence of the genus Homo: From concept to taxonomy

The main goal of this paper is to present an overview of hypotheses concerning early Homo specimens and to discuss the definition of the genus Homo in the light of recent discoveries. For some authors, all the specimens attributed to early Homo belong to one unique species. For others, this group (Homo habilis sensu lato) is heterogeneous and could be splitted into two groups: H. habilis and Homo rudolfensis. Some researchers have also proposed to put the species habilis and rudolfensis into the genera Australopithecus or Kenyanthropus. Therefore, two scenarios concerning first humans seem to emerge. An emergence of the genus Homo, as early as 2.8 Ma, with Homo sp. specimens and the species H. habilis and H. rudolfensis, another at 1.9 Ma with Homo ergaster. According to the recent archaeological and paleoanthropological discoveries, these criteria often considered to be crucial for the definition of the genus Homo, as the cranial capacity, the humanlike manipulative abilities, the habitual erect posture and bipedal gait, the language ability and the capacity to make tools are now obsolete.     

Découverte d’un nouvel hominidé à Dmanissi (Transcaucasie,Géorgie)

Four human remains: one mandible, two skulls and one metatarsus were discovered between 1991 and 1999 at the open-air site of Dmanisi, Georgia, in a precise stratigraphic, palaeontological and archaeological context, in volcanic ashes dated to 1.81 ± 0.05 Ma. The first studies of these fossils enable the authors to compare them with the morphology of archaic African Homo erectus, ascribed to Homo ergaster, and to ascertain hominid presence at the gates of Europe 300 000 years earlier than the classical scenario forecasted. In September 2000, the discovery of a second more complete and robust mandible D 2600 presents a threefold interest: palaeontological, functional and pathological. A comparison with Homo habilis and Homo erectus leads to the recognition of a new Homo species: H. georgicus sp. nov. The morphofunctional characteristics and the antiquity of H. georgicus characterise the root of a long Eurasian line.     

Homo floresiensis: a cladistic analysis

The announcement of a new species, Homo floresiensis, a primitive hominin that survived until relatively recent times is an enormous challenge to paradigms of human evolution. Until this announcement, the dominant paradigm stipulated that: 1) only more derived hominins had emerged from Africa, and 2) H. sapiens was the only hominin since the demise of Homo erectus and Homo neanderthalensis. Resistance to H. floresiensis has been intense, and debate centers on two sets of competing hypotheses: 1) that it is a primitive hominin, and 2) that it is a modern human, either a pygmoid form or a pathological individual. Despite a range of analytical techniques having been applied to the question, no resolution has been reached. Here, we use cladistic analysis, a tool that has not, until now, been applied to the problem, to establish the phylogenetic position of the species. Our results produce two equally parsimonious phylogenetic trees. The first suggests that H. floresiensis is an early hominin that emerged after Homo rudolfensis (1.86 Ma) but before H. habilis (1.66 Ma, or after 1.9 Ma if the earlier chronology for H. habilis is retained). The second tree indicates H. floresiensis branched after Homo habilis.     

Longgupo: EarlyHomo colonizer or late plioceneLufengpithecus survivor in south China?

A fragment of mandible and a maxillary incisor of different individuals from the Longgupo Cave, China have been cited as evidence of an early dispersal ofHomo from Africa to Asia. More specifically, these specimens are said to resemble “Homo ergaster” orHomo habilis, rather than the species usually thought to be the first Asian colonizer,Homo erectus. If this supposition is correct, it calls into question which hominid (sensu stricto) first left Africa, and why hominids became a colonizing species. Furthermore, the Longgupo remains have been used to buttress the argument thatHomo erectus evolved uniquely in Asia and was not involved in the origins of modern humans. We question this whole line of argument because the mandibular fragment cannot be distinguished from penecontemporary fossil apes, especially the Late Miocene-Pliocene Chinese genusLufengpithecus, while the incisor is indistinguishable from those of recent and living east Asian people and may be intrusive in the deposit. We believe that the Longgupo mandible represents the relic survival of a Late Miocene ape lineage into a period just prior to the dispersal of hominids into southeastern Asia, with some female dental features that parallel the hominid condition. If the Longgupo mandibular fragment represents a member of theLufengpithecus clade, it demonstrates that hominoids other thanGigantopithecus and the direct ancestor of the orangutan persisted in east Asia into the Late Pliocene, while all other Eurasian large-bodied hominoids disappeared in the Late Miocene.     

Dental microwear texture analysis of hominins recovered by the Olduvai Landscape Paleoanthropology Project, 1995-2007

Dental microwear analysis has proven to be a valuable tool for the reconstruction of aspects of diet in early hominins. That said, sample sizes for some groups are small, decreasing our confidence that results are representative of a given taxon and making it difficult to assess within-species variation. Here we present microwear texture data for several new specimens of Homo habilis and Paranthropus boisei from Olduvai Gorge, bringing sample sizes for these species in line with those published for most other early hominins. These data are added to those published to date, and microwear textures of the enlarged sample of H. habilis (n = 10) and P. boisei (n = 9) are compared with one another and with those of other early hominins. New results confirm that P. boisei does not have microwear patterns expected of a hard-object specialist. Further, the separate texture complexity analyses of early Homo species suggest that Homo erectus ate a broader range of foods, at least in terms of hardness, than did H. habilis, P. boisei, or the “gracile” australopiths studied. Finally, differences in scale of maximum complexity and perhaps textural fill volume between H. habilis and H. erectus are noted, suggesting further possible differences between these species in diet.     

Evolution of mandibular arcades: a statistical study

The alveolar arcades of a large number of fossil mandibles including Australopithecus and hominids fromHomo habilis andHomo erectus up to modern man have been characterized by fourteen cartesian points each representing a tooth. From these points, dimensions and angles have been calculated. These values are correlated to the geological age of the fossils. A linear dependance of dimensions and angles on the logarithm of age has been found. These results are discussed in the framework of a continuous gradual development within genus Homo and contrasted to prehominid data. Using these mean arcades and selected angles thereon the European and the AfroasiaticHomo erectus are compared and contrasted to the Neandertalians.     

The history of the genusHomo

The genusHomo was established by Carolus Linnaeus in 1758. During the course of the past 150 years, the addition of fossil species to the genusHomo has resulted in a genus that, according to the taxonomic interpretation, could span as much time as 2.5 Myr, and include as many as ten species. This paper reviews the fossil evidence for each of the species involved, and sets out the case for their inclusion inHomo. It suggests that while the case for the inclusion of some species in the genus (e.g.Homo erectus) is well-supported, in the case of two of the species,Homo habilis andHomo rudolfensis, the case for their inclusion is much weaker. Neither the cladistic evidence, nor evidence about adaptation suggest a particularly close relationship with laterHomo.     

Conclusion. Bilan des recherches interdisciplinaires effectuées sur le site de l’Homo erectus de Denizli,Kocabaş, Bassin de Denizli,Anatolie, Turquie. L’Homo erectus aux portes de l’Europe

The study of the site of Kocaba?, which yielded an archaic Homo erectus skullcap, was undertaken in 2011 and 2012, at the request of Professor Mehmet Cihat Alçiçek. This interdisciplinary French–Turkish research programme comprised the geochronological, magnetostratigraphic, biochronological and paleoenvironmental study of the site and the paleoanthropological study of the skullcap itself. The association of large mammals enabled us to attribute the travertine formations bearing the skullcap to the second part of the Upper Pleistocene, and more specifically to between 1.5 and 1.2 million years, because of the disappearance or appearance of certain species. This biochronological age is confirmed by the paleomagnetism study, which places the travertines bearing the skullcap in a period of reversed polarity, underlying a normal polarity formation, which could be attributed to the Cobb Mountain paleomagnetic excursion, dated to 1,194,000 years. The dating of these fauna by the ²⁶Al/¹⁰Be cosmogenic nuclide method by Anne-Elisabeth Lebatard yielded an age older than 1.22 Ma and more recent than 1.5 Ma. The Hominid skullcap from this formation can be attributed to a Homo erectus, slightly more evolved than those of Homo ergaster KNM-ER 3733 (1.78 Ma) and KNM-ER 15,000 (1.5 Ma), similar to that of Daka (Bouri), which is about a million years old and older than the Bodo fossil (estimated at 600,000 years) and Kabwe (between 300,000 and120,000 years). The archaic Homo erectus skullcap from Kocaba?, referred to as Denizli Man, proves that Homo erectus was already present in Anatolia, at the crossroads of Africa, Asia and Europe, a little more than 1.2 million years ago.     

L'hominidé du Pléistocène supérieur en Corée

Since the middle of 1970s, one rock-shelter, six caves and one open archaeological site have yielded several hominid fossils. Among them, fossils from Yonggok, Mandal, Sangsi and Hungsu cave sites are useful to reconstruct the general shape of the Upper Pleistocene Hominids in Korea. The main study of this paper is to focus on the reconstruction of the general shape of the upper Pleistocene hominids, and to examine the cause of cranial changes from the late upper Paleolithic to Mesolithic period in Korea. Biomechanical principles are also applied to reconstruct the social activity of the upper Paleolithic man in Korea. In addition, it is assumed that main proponent of Paleolithic cultures in Korea might be Homo sapiens rather than any other species. Based on the anatomical characteristics of skull, the axillary border of the scapular and the midshaft of the femur, general appearance of Homo sapiens during the Pleistocene in Korea could be reconstructed.     

Réflexions sur l’Acheuléen d’Anatolie

The Anatolia occupies one of the main routes for the dispersal of Homo erectus into the Eurasia. The Acheulean bifaces found on each region of Anatolia are the most important evidences of this situation. This vast distribution of the Acheulean bifaces in Anatolia indicates that all of the Anatolia should stay in the Movius Line. This means that the Movius Line should be reexaminate. Recently, the fossil remains of Homo erectus found in Dmanisi (Georgia) and their very old dates around 1.8 million years put forward the importance of Anatolia one more time. Homo erectus who came in Anatolia by following the Levant Corridor might used the Anatolian bridge for passing to the Transcaucasia. If the well-preserved cave site on the line that expands from Hatay to Kars in Anatolia founds and excavates, it will prove additional information some problems about the Homo erectus movements and distribution of Acheulean Industrial Tradition in West Asia. This paper reviews the evidence for the Acheulean in Anatolia and discusses the distribution of Acheulean bifaces in Anatolia, generally found in open air site.     

Homo erectus from the Yunxian and Nankin Chinese sites: Anthropological insights using 3D virtual imaging techniques

Recent applications of 3D virtual imaging techniques in human palaeontology have increased the possibilities and the accuracy of anthropological analysis. Two examples are given for the reconsideration of fossils discovered more than 20 years ago, thanks to this new technology. The Lower and Middle Pleistocene skulls from Yunxian and Nankin in China, which were damaged in the process of fossilization, have been virtually reconstructed. A detailed reinvestigation has been conducted by considering those reconstructed skulls and their unpublished characters (i.e., inner anatomical features inaccessible until now). The results of this analysis provide new information about the early hominids of China and contribute to the discussion of variability in Homo erectus.     

Les industries lithiques préoldowayennes du début du Pléistocène inférieur du site de Dmanissi en Géorgie

The lithic industry discovered at the Dmanissi site, in Georgia is dated to between 1.81 and 1.7 Myrs and is in association with a rich faunal assemblage composed of large Quaternary vertebrates, as well as several hominid fossils attributed to Homo georgicus, and attests to the human presence on the border of Europe at the beginning of the Lower Pleistocene. The material taken into account in this study was excavated from 1991 to 1999 and comprises 4446 lithic pieces coming from Beds I through VI of the site. The assemblage is very homogenous from the base to the top of the deposits and shows no significative evolutionary tendencies. The lithic material includes a high proportion of whole pebbles (33.8% of the assemblage) coming from two nearby rivers, the Mashavera and the Pinezaouri. They are essentially of fine and coarse grained volcanic tuff, basalt, but also of rhyolite, granite, quartz, as well as other volcanic and metamorphic rocks. Pebbles used for percussion, shaping or debitage were chosen according to their petrographic nature, their morphology and their size. Whole pebbles with percussion marks situated on their extremities or with isolated removals showing convexe edges, are abundant (1.3% of the assemblage). Other pebbles showing percussion marks on a flat face, were used as anvils. Broken pebbles and pebble fragments are very numerous (30.4% of the assemblage). These often show percussion marks on their cortical surfaces. Fractures are generally related to violent percussion as the pebbles were used for striking instruments, or as they were intentionally broken. Some fractures may have been caused accidentally during flaking. Pebble tools represent 4.8% of the lithic assemblage and 10% of the industry, excluding whole and fractured pebbles. These include essentially the primary choppers (pebbles with isolated concave removal negatives) (6% of the industry and 60.1% of the pebble tools), choppers showing continuous cutting edges without a point (2.1% of the industry and 21.2% of the pebble tools). Chopping-tools are very rare (0.8% of the industry and 8.7% of the pebble tools). Although choppers without pointed cutting edges were made using very few removals (3.3 on average), they usually present a regular cutting edge and seem relatively standardised. Cores are well represented (5% of the industry, excluding whole and broken pebbles). They are characterized by a low degree of exploitation and by a frequence of cortical striking platforms. Cored knapped on a single face are most frequent, representing nearly half of the pieces (42.3%), while bifacial cores are present in smaller proportions (34.2%) and multifacial cores are rare (6.3%). Non-modified flakes are very numerous and usually of small size and intentional retouch is absent. On the other hand, the cuttingedges of many the pieces; broken pebbles, pebble tools, cores and flakes, show irregular micro-retouch and irregular retouch such as isolated notches or with continuous or overlapping configuration, sometimes associated with localised crush marks which appear to have been caused by intensive use and heavy working of the pieces. A total of 31.3% of the non-modified flakes show irregular retouch on their cutting edges. One of the main characteristics of the Dmanissi industry appears therefore to be the obtaining of flakes, most often of small size, to be used without modification. The technological and typological characteristics of the lithic industry from Dmanissi allow to attribute the assemblage to a "Pre-Oldowayen" cultural horizon (Lumley de et al., 2004), characterized by the absence of small retouched tools, which appears in East Africa from 2.55 Myrs ago. This cultural horizon is present at the border of Europe, at Dmanissi, around 1.81 Myrs ago and in Western Europe, on the shores of the Mediterranean, at Barranco León about 1.3 Myrs ago and at Fuente Nueva 3 about 1.2 Myrs ago. The lithic industry from the Dmanissi site seems anterior to the Oldowan cultural horizon, characterized by the presence of standardized small retouched tools, which appears in East Africa around 1.8 Myrs ago and emerges in Mediterranean Europe around 800?000 years ago.     

A multivariate study of the Petralona skull

The Petralona skull from Greece has been generally accepted as an Upper Pleistocene variant of Neanderthal Man. It was included in a series of multivariate analyses performed by the author on cranial measurements of Pleistocene hominids. The results of the Mahalanobis D² analyses suggest that Petralona is cranially similar to the Neanderthals, but facially is distinct. Overall it resembles the Broken Hill and Djebel Ighoud skulls most closely but there are features of resemblance to Homo erectus material. The results of these analyses support recent suggestions that the Petralona material is of greater antiquity than the Upper Pleistocence.     

A SINGLE LINEAGE IN EARLY PLEISTOCENE HOMO: SIZE VARIATION CONTINUITY IN EARLY PLEISTOCENE HOMO CRANIA FROM EAST AFRICA AND GEORGIA

The relationship between Homo habilis and early African Homo erectus has been contentious because H. habilis was hypothesized to be an evolutionary stage between Australopithecus and H. erectus, more than a half‐century ago. Recent work re‐dating key African early Homo localities and the discovery of new fossils in East Africa and Georgia provide the opportunity for a productive re‐evaluation of this topic. Here, we test the hypothesis that the cranial sample from East Africa and Georgia represents a single evolutionary lineage of Homo spanning the approximately 1.9–1.5 Mya time period, consisting of specimens attributed to H. habilis and H. erectus. To address issues of small sample sizes in each time period, and uneven representation of cranial data, we developed a novel nonparametric randomization technique based on the variance in an index of pairwise difference from a broad set of fossil comparisons. We fail to reject the hypothesis of a single lineage this period by identifying a strong, time‐dependent pattern of variation throughout the sequence. These results suggest the need for a reappraisal of fossil evidence from other regions within this time period and highlight the critical nature of the Plio‐Pleistocene boundary for understanding the early evolution of the genus Homo.     

Variation among early Homo crania from Olduvai Gorge and the Koobi Fora region

Fossils recognized as early Homo were discovered first at Olduvai Gorge in 1959 and 1960. Teeth, skull parts and hand bones representing three individuals were found in Bed I, and more material followed from Bed I and lower Bed II. By 1964, L.S.B. Leakey, P.V. Tobias, and J.R. Napier were ready to name Homo habilis. But almost as soon as they had, there was confusion over the hypodigm of the new species. Tobias himself suggested that OH 13 resembles Homo erectus from Java, and he noted that OH 16 has teeth as large as those of Australopithecus. By the early 1970s, however, Tobias had put these thoughts behind him and returned to the opinion that all of the Olduvai remains are Homo habilis. At about this time, important discoveries began to flow from the Koobi Fora region in Kenya. To most observers, crania such as KNM-ER 1470 confirmed the presence of Homo in East Africa at an early date. Some of the other specimens were problematical. A.C. Walker and R.E. Leakey raised the possibility that larger skulls including KNM-ER 1470 differ significantly from smaller-brained, small-toothed individuals such as KNM-ER 1813. Other workers emphasized that there are differences of shape as well as size among the hominids from Koobi Fora. There is now substantial support for the view that in the Turkana and perhaps also in the Olduvai assemblages, there is more variation than would be expected among male and female conspecifics. One way to approach this question of sorting would be to compare all of the new fossils against the original material from Olduvai which was used to characterize Homo habilis in 1964. A problem is that the Olduvai remains are fragmentary, and none of them provides much information about vault form or facial structure. An alternative is to work first with the better crania, even if these are from other sites. I have elected to treat KNM-ER 1470 and KNM-ER 1813 as key individuals. Comparisons are based on discrete anatomy and measurements. Metric results are displayed with ratio diagrams, by which similarity in proportions for several skulls can be assessed in respect to a single specimen selected as a standard. Crania from Olduvai examined in this way are generally smaller than KNM-ER 1470, although OH 7 has a relatively long parietal. In the Koobi Fora assemblage, there is variation in brow thickness, frontal flattening and parietal shape relative to KNM-ER 1470. These comparisons are instructive, but vault proportions do not help much with the sorting process. Contrasts in the face are much more striking. Measurements treated in ratio diagrams show that both KNM-ER 1813 and OH 24 have relatively short faces with low cheek bones, small orbits and low nasal openings. Also, they display more projection of the midfacial region, just below the nose. This is not readily interpreted to be a female characteristic, since in most hominoid primates the females tend to have flatter lower faces than the males. The obvious size differences among these individuals have usually been interpreted as sex dimorphism, but, in fact, two taxa may be sampled at Olduvai and in the Turkana basin at the beginning of the Pleistocene. One large-brained group made up of KNM-ER 1470, several other Koobi Fora specimens, and probably OH 7, can be called Homo habilis. If these skulls go with femora such as KNM-ER 1481 and the KNM-ER3228 hip, then this species is close in postcranial anatomy to Homo erectus. The other taxon, including small-brained individuals such as KNM-ER 1813 and probably OH 13, seems also to be Homo rather than Australopithecus. If the OH 62 skeleton is part of this assemblage, then the small hominids have postcranial proportions unlike those of Homo erectus. However, it is too early to point unequivocally to one or the other of these groups as the ancestors of later humans. Both differ from Homo erectus in important ways, and both need to be better understood before we can map the earliest history of the Homo clade. © 1993 Wiley-Liss, Inc.     

The Ceprano Skull and the Earliest Peopling of Europe: Overview and Perspectives

This paper deals with the cranial remains belonging to the earliest hominids found in Europe over the past 15 years. Morphological and metric traits were scored and compared with the data on the pleistocenic remains found in the Old World. The cladistical and multivariate analyses – carried out respectively on the morphological and metric traits – seem to suggest that the Asian remains attributed to Homo erectus differ significantly from those found in the western territories (southern Africa and northern Europe). Among such reports, further differences between the earliest African forms (H. habilis and H. ergaster) and the most recent European and African H. heidelbergenis are worth mentioning. Relying upon this research, the Ceprano calvaria seems to represent a new species (H. cepranensis) from which the later forms – specifically H. heidelbergenis – originated. The presence of some heidelbergensis traits observed in nuce on the Ceprano specimen seems to support this scenario.