Immunospecificity of albumin detected in 1.6 million-year-old fossils from Venta Micena in Orce,Granada, Spain

The Orce skull fragment from southern Spain, dated at 1.6 Myr, has been a subject of heated controversy since it was first discovered in 1982. If it is hominid, as its discoverers contend, it is by far the oldest fossil hominid yet found in western Europe and implies that human populations settled this region much earlier than was previously realized. Numerous stone artifacts found at the Orce sites provide evidence that hominids were indeed present there in the Lower Pleistocene. Some paleontologists maintain that the 8 cm diameter occipital fragment is from a horse, not a hominid. Two independent investigations of the residual proteins in the skull were undertaken, one at the University of Granada in Spain, the other at the University of California, San Francisco. Two immunological methods of comparable sensitivity were employed for detection and species attribution of protein extracted from fossil bone: the Granada team used an enzyme-linked-immunosorbent assay (ELISA), and the UCSF team used a radioimmunoassay (RIA). Both teams obtained reactions characteristic of human albumin in the Orce skull and horse albumin in some of the horse fossils. These results support the lithic evidence that hominids were living in Andalusia 1.6 million years ago. Am J Phys Anthropol 103:433–441, 1997. © 1997 Wiley-Liss, Inc.     

Réévaluation de la présence humaine au Pléistocène inférieur dans le Sud de l'Espagne

The Early Pleistocene locality at Venta Micena (Orce, Guadix-Baza basin, province of Granada, Spain) has provided four fossil remains - skull fragment VM-0, and long bone diaphyses VM-1960, VM-3691, and VM-12000 - which have been tentatively attributed to the hominids. Although several methodologies have been used to ascertain the human affinities of these specimens - including anatomical, morphometric and immunological analyses - the results obtained have not been conclusive, instigating a persistent debate. A taphonomic approach is used here for estimating the probability that a taxon the size of Homo sp. (~ 50 kg) could be represented in the fossil assemblage by four bone fragments and no tooth remain. A least-squares regression analysis between the percentage of teeth and the body mass estimated for each taxon of large mammals (N = 20) predicts a raw abundance of six teeth for Homo sp. in the assemblage. Given that up to the present moment no tooth remains attributable to the hominids has been unearthed during systematic excavations in the Venta Micena quarry, which has provided more than 15,000 fossils of large mammals, this argues strongly against the possibility that the three bone specimens could belong to Homo sp. The phalanx CV-0 from the Early Pleistocene site of Cueva Victoria (Cartagena, Spain) has also been attributed to the genus Homo. The taxonomic assignment of this specimen is biased, however, because it was not compared with Theropithecus oswaldi, the only primate species actually recorded from this karstic locality. A comparative anatomical and morphometric analysis of fossil and modern specimens of Theropithecus suggests that CV-0 can be attributed to T. oswaldi. As a result, Cueva Victoria does not contribute additional information concerning the first human settlements in Europe. By these reasons, apart from the paleoanthropological and archaeological findings from Atapuerca (TD lower levels and Sima del Elefante), the rich archaeological assemblages from Barranco León and Fuente Nueva-3 in Orce, dated 1.3-1.2 Myrs, which include fourteen hundred stone tools of Oldowan technology, constitute at present the only unequivocal evidence of human presence in Southeast Spain during Early Pleistocene times.     

Some comments on the case for Early Pleistocene hominids in South-Eastern Spain

Several sites in the Orce Basin have revealed evidence of the presence of hominids in the Early Pleistocene. These remains are dated to over 1.0 million years, while they may be as old as 1.6 million years. The skeletal remains from Venta Micena in the Orce Basin show a molecular “fossil protein” pattern which aligns them with hominids, but not with equids. This is supported by the anatomical evidence of the two humeral shafts from theEstrato Blanco in the Venta Micena deposits. The biparieto-occipital partial calvaria shows some unusual features if VM-0 is a hominid specimen. the presence of a prominent crest on the internal surface of the occipital fragment adjacent to the point lambda is decidedly unusual for a modern human calvaria. Moreover, theimpressions gyrorum, in the region where the superior parietal lobule of the cerebral hemisphere abutted against the calvaria, point to a bipartite superior parietal lobule with anterior and posterior moieties which, on the endocast, are clearly separated by a depression that represents a sulcus. These morphological traits are rather puzzling if VM-0 is a hominid, and at first they led me to hesitate over the anatomical identification of VM-0. However, the studies of Campillo (1989) and of Campillo and Barcelo (1986) suggest that the features of the fragment VM-0 are compatible with those of a hominid. Because I believe that we do not possess sufficient information on the variability of the endocranial and ectocranial manifestations of the sagittal suture and of its variance with age of the individual, in different hominid species and different equid species, I have not adduced this pattern as evidence in support or rebuttal of the hominid status of VM-0.     

Fractal analysis of the Orce skull sutures

Methods of fractal geometry (Mandelbrot, 1983) are used here to analyse the relative complexity of the sagittal and lambdoid sutures visible in the skull fragment formed by parts of an occipital squame and parietals found in a sealed deposit at the early Lower Pleistocene site of Venta Micena (Orce, Granada, Spain), generally regarded as human bone but occasionally suggested as belonging to an equid. For comparison with the fossil, corresponding sutures of various primates (hominids, pongids and cercopithecids) and two other groups of mammals (equids and ruminants) were analysed using the computer program FRACTAL-D (Slice, 1989) in order to determine their fractal dimensions as a measure of differential sutural design complexity. The results show that the fractal dimension of the Venta Micena skull sutures lies within the range of variation for infant specimens of both modern and Plio-Pleistocene hominids. Sutural complexity in young pongids and cercopithecids overlaps the range of fractal dimensions found in hominids, whereas values obtained from equids and ruminants are significantly greater than those for all the primates analysed here. Therefore, in terms of fractal dimension measures of relative complexity, the sutures preserved in the Venta Micena fossil could not have belonged to an equid (pace Agusti & Moyà-Sola, 1987); rather, its fractal dimension is consistent with the attribution of the fossil to an infant of Homo sp.     

Dispersion du Genre Homo au sud d’Ibérie et au Maghreb. Nouvelles données à propos des fossiles VM-1960 et BL-0

In this article, there is a discussion about out-of-Africa models of Homo dispersal, considering new findings, specially a new site with Oldowan industry in Morocco (Ain-Beni-Mathar), found in 2005 by some of the authors (Gibert J, Gibert L, and El Hamouti). There is also a critical of the models of Homo dispersal only based on carrion consumption. We also discuss some objections about two human remains from Orce sites (VM-1960 and BL-0) based on morphometric data (that, in any case, move these remains away from genus Homo), on authority criteria and ad hoc hypothesis, ignoring the decisive anatomical data.     

Length estimate for KNM-ER 736, a hominid femur from the lower pleistocene of East Africa

Our knowledge concerning stature in earlyHomo is scanty. In this paper, based on comparison with the fossil femur KNM-ER 999, an estimate of 482 mm femur length is derived for KNM-ER 736, the latter dating from the Lower Pleistocene. From comparison with other fossil and modern femora, KNM-ER 736 appears to be the longest hominid femur so far recovered from a site of Early Pleistocene age. Moreover, the estimated femur length is higher than the published mean values of most modern populations. Provided that trunk and head proportions were not radically different from modernH. sapiens, the finding would suggest that a stature similar to that of modern man was already reached by East AfricanHomo as early as about 1.6 Myr before present.     

Anatomical study: comparison of the cranial fragment from venta micena, (orce; Spain) with fossil and extant mammals

The fragment of the skull of Orce attributed to the genusHomo is compared with fossil and extant mammals. The anatomical analysis supports the idea of ascribing it to an infantile individual of the genusHomo, close to the primitive Turkana specimens.     

Spanish late Pliocene and early Pleistocene hominid, palaeolithic and faunal finds from Orce (Granada) and Cueva Victoria (Murcia)

In SE Spain, recent excavations in the Orce basin and at Cueva Victoria indicate presence of both hominids and hominid activity from the Plio-Pleistocene boundary and early Lower Pleistocene.     

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.     

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.     

Model of a hypothetical protohominid dentition and its implications for hominid phylogeny

The complete dentition of the common ancestor ofAustralopithecus andHomo, intermediate between that of a pongid and a hominid, is virtually unknown. The maxillary dentition (P3-M2) ofRamapithecus brevirostris Lewis, 1934, a pongid from the Early Pliocene, and that of hominids from the Late Pliocene and Plio/Pleistocene is known. SinceR. brevirostris is probably ancestral to the hominids, a model of intermediate maxillary dentition (P3-M2) is extrapolated and described. The model represents a hypothetical protohominid dentition. It does not conform with the teeth ofAustralopithecus, but shows greater morphological affinity to hominine dentition and to 5 myo hominids. TheHomo lineage, therefore, may go back to the Middle Pliocene. According to the normal sequence of evolution, it is most unlikely thatAustralopithecus gave rise toHomo, but much more probable that a very early, generalizedHomo evolved into an advanced, specializedAustralopithecus.     

Radiographical study of skull fragment of Venta Micena (VM-0) (Orce, Granada Spain)

Since 1983, the Orce skull has been a polemic piece. The last article which refuses the human affinity of VM-0 is based on the presence of a supposed coronalis suture in VM-0 situated at 40mm of lambda point (Moyà-Solà &; Khöler 1997). This would imply that VM-0 belonged to a very small skull not possible in human genus. The present detailed x-ray study which includes: “classic x-ray”, digital x-ray, macro x-ray and computerized tomography, proves definitively the not presence of coronalis suture in VM-0 and demonstrate that all interpretations based on the presence of coronalis suture are a great mistake. Also this study shows different characters which prove the human affinity of VM-0: presence of a wide sulcus sagittalis (9mm), the print of the superior sinus sagittalis where drained venous blood of a big brain is not possible in a colt whose brain size is similar to that of sheep. We consider that the visual inspection is not enough in human Palaeontology and it is incorrect to not apply the different tests which permit proving or refusing the different subjective impressions.     

Nouvelles découvertes de dents d’hominidés dans le membre Kaitio de la formation de Nachukui (1,65–1,9 Ma), Ouest du lac Turkana (Kenya)

New hominid teeth from the Kaitio member (1.65–1.9 Myr) in West Turkana (Kenya). New hominid teeth have been recovered from the archaeological sites of Kokiselei 1 and Naiyena Engol 1. These two sites are located in the west side of the Turkana Basin and belong to the Kaitio member of the Nachukui Formation. They are dated between 1.65-1.79 and 1.7-1.8 Myr respectively. The four teeth (left maxillary canine and first molar, right maxillary third molar and left mandibular third molar) discovered in Kokiselei 1 are attributed to Australopithecus boisei. The right mandibular first premolar found in Naiyena Engol 1 is referred to Homo sp. aff. ergaster. To cite this article: S. Prat et al., C. R. Palevol 2 (2003).     

The “Phoca standard”: An external molecular reference for calibrating recent evolutionary divergences

Comparison of the complete mitochondrial DNA (mtDNA) of the high-Arctic ringed seal (Phoca hispida) and the sub-Arctic harbour (P. vitulina) and grey (Halichoerus grypus) seals shows that they are genetically equidistant from one another. We relate the evolutionary divergence of the three species to expanding glaciation in the Arctic Basin and establish, in conjunction with mtDNA data, a standard reference for calibration of recent divergence events among mammalian taxa. In the present study, we apply the “Phoca standard” to the dating of divergences within the hominid phylogenetic tree. After determining the relative rates of substitution over all mitochondrial protein-coding genes in the different evolutionary lineages, we estimate that humans and chimpanzees diverged from each other 6.1 Mya (95% confidence limits: 5.2–6.9 Mya). The corresponding lower-limit divergence between common chimpanzee,Pan troglodytes, and pygmy chimpanzee,P. paniscus, occurred 3 (2.4–3.6) Mya, and the primary split within theP. troglodytes complex 1.6 (1.3–2.0) Mya. The analyses suggest that the split betweenGorilla andPan/Homo occurred 8.4 (7.3–9.4) Mya. They also suggest thatPongo (orangutan) and the lineage leading to gorillas, chimpanzees, and humans diverged 18.1 (16.5–19.6) Mya. The present analysis is independent of the hominid paleontological record and inferential morphological interpretations and thus is a novel approach to the lower-limit dating of recent divergences. Correspondence to: U. Arnason     

The first human dispersal to Europe: Remarks on the archaeological and palaeoanthropological record from Orce (Guadix-Baza basin, southeastern Spain)

The chronology of the first arrival ofHomo in Europe is a rather controversial issue, with most scholars claiming until very recently that there were no permanent human settlements before the middle Pleistocene. However, new findings at Atapuerca, Dmanisi and Orce, as well as the re-evaluation of the evidence from Java, Israel and China indicate a protracted chronology for the arrival of hominids in Eurasia, during late Pliocene/lower Pleistocene times. The systematic study of the macrovertebrate assemblages from circummediterranean sites such as Orce and Dmanisi has shown a faunal replacement at the Plio-Pleistocene boundary, marked by the arrival in Europe of African immigrants such as the hippopotamus, an equid similar to modern grevy’s zebra, a large cercopithecoid and several carnivores, including a giant hyaena, a sabre-tooth and a wild dog. An analysis of the relative frequency of bones and teeth for those species of large mammals preserved in Venta Micena indicates the improbability thatHomo sp. is represented in the fossil assemblage by several bones and no tooth remain. Finally, new data on the stratigraphy of Barranco León are offered, with remarks on the discovery in this locality of lithic artefacts and molar tooth fragment BL5-0.     

A new species of the genusHomo (primates: Hominidae) from the Plio/Pleistocene of Koobi Fora,in Kenya

Reassessment of the hominine cranium, KNM-ER 1813, from the Plio/Pleistocene of Koobi Fora, in Kenya, shows that it is not a small-brained, extreme female variant ofH. habilis Leakey, Tobias, & Napier, 1964. Its cranial and dental morphology, morphometrics, and proportions do not conform with eitherH. habilis orH. antiquus Ferguson, 1984. On the basis of its distinctive morphological pattern and mensural gaps which distinguish it fromH. habilis andH. antiquus, the cranium KNM-ER 1813 is described as a common variant representing a male of a small-brained, intermediate population linkingH. habilis 1.83 Myr BP withH. antiquus 2.9 Myr BP, and a new paleospecies of the genusHomo. A key to the Homininae is provided and the phylogenetic relationship of KNM-ER 1813 toH. habilis andH. antiquus is discussed. This paper is dedicated to the memory of my wife,Grace, whose assistance will be sorely missed.     

The influence of mass,volume and density on the frequency of recovery of fossil hominid hand and wrist bones

A study of the mass, volume and density of each of the wrist and hand bones of male and female human skeletons was undertaken. It was found that the mass and volume (i.e. size) of the bones are well correlated with the relative frequencies of preservation ofAustralopithecus and earlyHomo wrist and hand bones from fossil hominid sites in Africa. In general, the larger the bone, the greater its preservation frequency. In contrast to findings on bovid bones, the density of hand and wrist bones is not well correlated with the frequency of such bones recovered from these sites. These findings may be explained in terms of the agents of deposition of the bones, the physical nature of the deposit, and the methods of extraction of the fossils from the deposit.     

Insights into the evolution of child growth from Lower Pleistocene humeri at Venta Micena (Orce, Granada province, Spain)

Well-defined human anatomical characteristics are present on humeral fragments of a child (VM-1960) and an adult (VM-3691) from early Lower Pleistocene sediments at Venta Micena: both have narrower medullary cavities than in AfricanHomo erectus/ergaster (KNM-ER 1808), and the child’s humeral shaft is longer than in recent 8-to-9-year-olds even though its muscle markings are less pronounced than theirs. We infer that exposure of growing children to high mechanical loading favoured Plio-Pleistocene skeletal evolution inHomo of humeral robusticity and elongation. Precocious childhood arm-bone development, occurring before pubertal growth-spurt increments in shoulder and arm muscularity, implies a different balance from today between prepubertal hormonal influences exerted on ossification (growth hormone and somatomedin C) and the adolescent gonadal hormones of our modern growth spurt which may have still been in the process of evolution by natural selection.     

Serial allocations of isolated mandibular molars of unknown taxonomic affinities from the Shungura and Unso Formations,Ethiopia, a combined method approach

The early hominid dental remains from the Omo succession represent a fragmentary but important source of information regarding hominid evolution during the 2 to 3 myr time period. As an initial step toward the evaluation of taxonomic affinities and evolutionary significance, the present study attempts serial allocations of 21 isolated mandibular molars from the Shungura and Usno Formations. A comparative sample consisting of 250 mandibular molars ofA.afarensis, A.africanus, A.robustus, A.boisei and earlyHomo was used to compile the baseline data for allocating the isolated Omo molars to serial positions. The methods employed in the present study include morphometric analyses of 5 cusp areas, 8 linear variables reflecting crown shape, and 4 measurements of fissure pattern. It was found that by combining morphological observations with both “restricted” and “non-restricted” applications of discriminant function analyses (sensu Albrecht, 1992), sufficiently reliable serial allocations could be attained.     

Plio-Pleistocene cercopithecids from Kanam East, western Kenya

This paper describes cercopithecid craniodental and postcranial fossils recovered by L. S. B. Leakey at Kanam East, Kenya during the early 1930s. These fossil monkeys have been generally assumed to have been derived from early Pliocene horizons, but their exact geographical and stratigraphical provenience is unknown. Although the question of the evolutionary significance of these specimens must await the recovery of more securely dated material from Kanam East, some general conclusions can be drawn concerning their taxonomic affinities and paleobiology. Based on comparative studies of the craniodental material, at least three extant genera are represented—Colobus,Lophocebus, andCercopithecus. The postcranial fossils include a number of hindlimb specimens, as well as the manubrium of a sternum and a caudal vertebra. Identification of the postcranial remains to particular genera is not possible, but they are similar in morphology to modern arboreal and semiterrestrial cercopithecid monkeys of small to medium size. It is evident that Kanam East had a diverse cercopithecid community, similar to those found today in forested and woodland habitats, and this may be of some significance in reconstructing the paleoecology of the site. Because the fossil record of most extant cercopithecid genera is rather sparse at Plio-Pleistocene sites in Africa, Kanam East represents one of only a few sites that has yielded material that can be assigned toColobus,Lophocebus, orCercopithecus. The fossil monkeys from the site, therefore, provide additional evidence to help reconstruct the paleobiology, as well as the patterns of species diversity and community structure that characterized the cercopithecid radiation during the Plio-Pleistocene.