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.     

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.     

Out of Africa and back again

If the genusHomo did indeed originate in Africa, then it must have spread by about 2 m.y. ago into Asia where it is represented at 1.8 m.y. ago byHomo erectus fossils. This latter species in turn eventually spread back into Africa, as indicated by the 1.4 m.y. old OH 9 calvaria from Olduvai, and into Europe, as indicated by the 800,000 year old Ceprano calvaria from Italy. These hominids are associated only with Oldowan style artefacts of cores, choppers and flakes and were apparently not conversant with Acheulean handaxe technology. It seems that they most probably evolved viaHomo heidelbergensis into the Neanderthals. Meanwhile, a completely separate development originating withHomo ergaster of about 1.7 m.y. ago in Africa and possessing Acheulean handaxe technology evolved via such forms as Ndutu and Steinheim intoHomo sapiens.     

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.     

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.     

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.     

Early Hominid diversity, age and biogeography of the Malawi-Rift

Remains of earlyHomo andParanthropus have been recovered from two contemporaneous sites (Uraha and Malema) in the “Hominid Corridor” in Northern Malawi (Chiwondo Beds). Faunal dating suggests an age of 2.5–2.3 Ma for both hominids. The two specimens, a mandible attributed toHomo rudolfensis (UR 501 from Uraha), and a maxillary fragment ofParanthropus boisci. (RC 911 from Malema) known only from eastern Africa, represent the southernmost known distribution of these taxa. The biogeographic significance of these hominids from the Malawi-Rift lay in their association with the eastern African endemic animal group. Biogeographic variation in south-eastern Africa may be linked to habitat change occurring due to climate change, with maximum change occurring around 2.5 Ma.     

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.     

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.     

巫山龙骨坡似人下颌属于猿类

本文将巫山似人下颌及其牙齿与东非早更新世人属 ,Dmanisi直立人以及元谋的禄丰古猿等做了比较。结果发现巫山标本的尺寸比前二者都小得多 ,而与后者很相符。巫山标本被有些人作为归属人属根据的那些特征大多是人和猿共具者 ,其中前臼齿齿根分叉则是在人类罕见 ,却是禄丰古猿的特征之一 ,前臼齿前接触面位置和跟座比例则反映猿类特征。最接近巫山下颌者是禄丰古猿 ,其间有否祖裔关系尚待更多标本来论证。     

Palaeopathology of the Pleistocene specimen D2600 from Dmanisi (Republic of Georgia)

Here we present a detailed palaeopathological study of the hominin mandible D2600 recovered at the Dmanisi site, Republic of Georgia. The Dmanisi assemblage represents the earliest evidence of hominins outside Africa with an age of 1.8 Ma. D2600 is the holotype of Homo georgicus species and its taxonomic assignment is still under debate. Our study reveals severe and unusual dental wear accompanied of extensive root exposure and dental axial migration, periapical abscesses and enamel fractures. In addition, there is evidence of post-eruptive tooth rotation and temporomandibular arthropathy. We propose that the wear pattern observed in this individual is related to a diet with a high intake of fibrous and abrasive foods such as fruits and plants, as it is usually recorded in chimpanzees and gorillas and unlike the wear pattern observed in other Homo specimens of our comparative sample. The rounded occlusal surfaces and highly polished labio-lingual surfaces of D2600 anterior teeth could be mainly the consequence of pre- and/or para-masticatory activities such as gripping and stripping. This type of food would be also the origin of the highly cupped occlusal morphology of the posterior dentition in combination with relatively slight approximal attrition. However, the lesions exhibited by D2600 have not significantly altered the morphology of the mandible and do not prevent a proper taxonomic assessment.     

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.     

Another nail in the coffin of the multiple-origins theory?

While mitochondrial sequences can be used to probe the time and place of the mitochondrial ‘Eve,’ nuclear genes can be used to ask a slightly different question: when did humans (members of the genus Homo) or their hominid precursors (the hominids) first leave Africa and fan out over Asia and Europe? If they did so recently, it seems likely that there was a recent African origin of our species, Homo sapiens, rather than multiple origins in various parts of the Old World. A recent paper⁽¹⁾ uses minisatellite data to make the argument that the departure from Africa happened very recently indeed. An alternative explanation for the data is that there was no single and irreversible departure from Africa, but that some peoples migrated back and forth between Africa and the rest of the Old World over the last few tens of thousands of years. For this and other reasons, putting a single date on the farewell to Africa remains problematical.     

The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

Liang Bua Homo floresiensis mandibles and mandibular teeth: a contribution to the comparative morphology of a new hominin species

In 2004, a new hominin species, Homo floresiensis, was described from Late Pleistocene cave deposits at Liang Bua, Flores. H. floresiensis was remarkable for its small body-size, endocranial volume in the chimpanzee range, limb proportions and skeletal robusticity similar to Pliocene Australopithecus, and a skeletal morphology with a distinctive combination of symplesiomorphic, derived, and unique traits. Critics of H. floresiensis as a novel species have argued that the Pleistocene skeletons from Liang Bua either fall within the range of living Australomelanesians, exhibit the attributes of growth disorders found in modern humans, or a combination of both. Here we describe the morphology of the LB1, LB2, and LB6 mandibles and mandibular teeth from Liang Bua. Morphological and metrical comparisons of the mandibles demonstrate that they share a distinctive suite of traits that place them outside both the H. sapiens and H. erectus ranges of variation. While having the derived molar size of later Homo, the symphyseal, corpus, ramus, and premolar morphologies share similarities with both Australopithecus and early Homo. When the mandibles are considered with the existing evidence for cranial and postcranial anatomy, limb proportions, and the functional anatomy of the wrist and shoulder, they are in many respects closer to African early Homo or Australopithecus than to later Homo. Taken together, this evidence suggests that the ancestors of H. floresiensis left Africa before the evolution of H. erectus, as defined by the Dmanisi and East African evidence.     

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.     

The environmental contexts of early human occupation of Georgia (Transcaucasia)

The hominid mandible and a third metatarsal found in Dmanisi (Republic of Georgia) are accompanied by a rich faunal assemblage and a core-chopper stone tool industry. The mandible represents a somewhat isolated morphological type of Homo erectus that appears, given the combination of its primitive and advanced traits and specific dental morphology, to be a forerunner of both late H. erectus and early archaic H. sapiens. The faunal assemblage mostly consists of Villafranchian mammals, with the majority of the species assigned to an early phase of the Upper Villafranchian (Late Villanian and Early Biharian). Faunal and paleobotanical evidence as well as the depositional nature of the site indicate that hominid occupation took place in a mosaic environment of open steppe and gallery forests. Both the concentration of resources and the warm climatic conditions in the Dmanisi region at the beginning of the early Pleistocene were favorable for hominid occupation. It is possible that hominids reached the Caucasus through the Levantine corridor, and that the environment of this region allowed them to establish a stronghold and later colonize adjacent areas.     

Dental microwear texture analysis and diet in the Dmanisi hominins

Reconstructions of foraging behavior and diet are central to our understanding of fossil hominin ecology and evolution. Current hypotheses for the evolution of the genus Homo invoke a change in foraging behavior to include higher quality foods. Recent microwear texture analyses of fossil hominin teeth have suggested that the evolution of Homo erectus may have been marked by a transition to a more variable diet. In this study, we used microwear texture analysis to examine the occlusal surface of 2 molars from Dmanisi, a 1.8 million year old fossil hominin site in the Republic of Georgia. The Dmanisi molars were characterized by a moderate degree of surface complexity (Asfc), low textural fill volume (Tfv), and a relatively low scale of maximum complexity (Smc), similar to specimens of early African H. erectus. While caution must be used in drawing conclusions from this small sample (n = 2), these results are consistent with continuity in diet as H. erectus expanded into Eurasia.     

The changing face of genus Homo

Since its introduction in the 18th century, the genus Homo has undergone a number of reinterpretations, all of which have served to make it a more inclusive taxon. In this paper, we trace this trend towards greater inclusiveness, and explain how it has affected the way Homo is defined. We then demonstrate that the current criteria for identifying species of Homo are difficult, if not impossible, to operate using paleoanthropological evidence. We discuss alternative, verifiable, criteria, and show that when these new criteria are applied to Homo, two species, Homo habilis and Homo rudolfensis, fail to meet them. We contend that the lowest boundary of Homo should be redrawn so that the earliest species in the genus is Homo ergaster, or early African Homo erectus. The appearance of this species at around 1.9 Myr appears to mark a distinct shift in hominin adaptive strategy involving morphological and behavioral innovations.