Species-specific albumin in fossil bones from Orce, Granada, Spain

A skull fragment (VM-0) from Orce, Granada, Spain, dated palaeomagnetically at about 1.6 Myr, is thought by some palaeontologist to be hominid, while others maintain it is equid. If hominid, it would be by far the oldest evidence ofHomo in Europe. Immunological studies on residual albumin in this fossil were carried out independently, and with different immunological methods, at the University of California, San Francisco (radioimmunoassay), and at the University of Granada, Spain (enzyme immunoassay). Other fossils attributed to hominids also studied wereVM1960 from Venta Micena, andCV-1 from Cueva Victoria, Murcia, Spain. Undisputed equid and bovid fossils from the same deposits and dated to a similar period as the Orce skull were also analyzed. Our results showed that species-specific albumin can be detected in 1.6 Myr-old hominid, equid and bovid fossils. The albumin from the Orce skull fragment and fromVM-1960 was immunologically closer to human albumin. These findings support the contention that theVM-0 andVM-1960 are hominid and that members of the genusHomo occupied southern Spain 1.6 Myr ago.     

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 and its applications in human palaentology

Fractal analysis was applied in human palaentology by Gibert and Palmqvist to estimate the value of the fractal dimension obtained from the cranial sutures preserved in the fragment of occipital bone (VM-0) found at the Venta Micena site. This paper also estimated the values of fractal dimension for different specimens in order to establish a taxonomy. Although that initial study demonstrated that the technique could be useful in human palaentology, the large variability of sutures observed in the VM-0 sample requires a mechanism that makes it possible to automatically obtain an objective plot of the suture to be analyzed.     

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.     

Evolution of femur and tibia in higher primates: Adaptive morphological patterns and phylogenetic diversity

Seventy six metrical traits measured on the femur and tibia of three higher primate groups —Ceboidea, Cercopithecoidea, Hominoidea have been processed by various univariate and multivariate statistical methods to survey the process of evolution of the morphology of the femur and tibia in higher primates. Intragroup and intergroup variability, similarity and differences as well as various aspects of scaling and sexual dimorphism have been analyzed to study adaptive trends and phylogenetic diversity in higher primates, in individual superfamilies and to explore the adaptive morphological pattern of early hominids and basic differences between hominids and pongids. Two basic morphotypes of the femur and tibia in higher primates have been determined. They are (1) advanced hominoid morphotype (hominids and pongids) and (2) ancestral higher primate morphotype (platyrrhine and cattarrhine monkeys, early hominoids, and hylobatids). Cebid lower limb bones are adapted to arboreal quadrupedalism with antipronograde features while femur and tibia of cercopithecid monkeys are basically adapted to the semi-arboreal locomotion. Early hominoids (Proconsul) and hylobatids are morphologically different from pongids; some features are close toAteles or other monkey species. Pongids and hominids are taken as one major morphological group with different scaling and some functional and morphological similarities. Numerous analogous features were described on the lower limb skeleton ofPan andPongo showing analogous ecological parameters in their evolution. Major morphological and biomechanical trends are analyzed. It is argued that early advanced hominoid morphology is ancestral both to the pongids and to early hominids. The progressive morphological trend in early hominids has been found fromA. afarensis with ancestral hominid morphology, toH. habilis with an elongated femur and structural features similar to advanced hominids. A detailed phylogenetic analysis of higher primate femur and tibia is also presented.     

Postcrania and the specific mate recognition system

The Recognition Concept of Species is examined for its potential usefulness in discriminating speciation events in the hominid fossil record. Controversies over species-specific characteristics amongHomo erectus and archaicHomo sapiens have centred on traits of the skull, largely because this element is most commonly preserved. Modern humans have an intuitive knowledge of their own Specific Mate Recognition System (SMRS), and therefore have the opportunity to compare their own SMRS to that of fossil hominids and the extent pongids. Such comparison suggests that our own skeletal SMRS may depend less on features of the skull than on the morphology of the postcranial anatomy. We propose that these components be further examined in this regard. We tentatively conclude that examination of the Recognition Concept of Species indicates that from lateHomo erectus onwards, the same SMRS has been shared in common by all hominids, including modernHomo sapiens. This suggests that, following the SMRS criterion, none of these forms can be categorized as separate species.     

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.     

Multivariate discriminant analysis of dental data bearing on early hominid affinities

The dimensions of hominoid dentitions are compared by multiple discriminant analysis. By this technique the fossil taxa are compared with living pongids and modern man in a multivariate framework. This enables the classification of the fossils to be made consistent with that of the living forms. H. africanus and H. erectus generally form the most compact grouping within the hominids, thus suporting the argument that these two species can indeed be lumped into a single genus. The degree of separation between H. africanus and Paranthropus is found to be at least as great as that between the genera of modern apes. Gigantopithecus sorts with the hominids rather than with the pongids and seems to be most closely related to Paranthropus.     

Paleoenvironment of Dryopithecus brancoi at Rudabánya, Hungary: evidence from dental meso- and micro-wear analyses of large vegetarian mammals

The environment of the hominoid Dryopithecus brancoi at Rudabánya (Late Miocene of Hungary) is reconstructed here using the dietary traits of fossil ruminants and equids. Two independent approaches, dental micro- and meso-wear analyses, are applied to a sample of 73 specimens representing three ruminants: Miotragocerus sp. (Bovidae), Lucentia aff. pierensis (Cervidae), Micromeryx flourensianus (Moschidae), and one equid, Hippotherium intrans (Equidae). The combination of meso- and micro-wear signatures provides both long- and short-term dietary signals, and through comparisons with extant species, the feeding styles of the fossil species are reconstructed. Both approaches categorize the cervid as an intermediate feeder engaged in both browsing and grazing. The bovid Miotragocerus sp. is depicted as a traditional browser. Although the dental meso-wear pattern of the moschid has affinities with intermediate feeders, its dental micro-wear pattern also indicates significant intake of fruits and seeds. Hippotherium intrans was not a grazer and its dental micro-wear pattern significantly differs from that of living browsers, which may suggest that the fossil equid was engaged both in grazing and browsing. However, the lack of extant equids which are pure browsers prevents any definitive judgment on the feeding habits of Hippotherium. Based on these dietary findings, the Rudabánya paleoenvironment is reconstructed as a dense forest. The presence of two intermediate feeders indicates some clearings within this forest; however the absence of grazers suggests that these clearings were most likely confined. To demonstrate the ecological diversity among the late Miocene hominoids in Europe, the diet and habitat of Dryopithecus brancoi and Ouranopithecus macedoniensis (Greece) are compared.     

Brain endocast asymmetry in pongids and hominids: Some preliminary findings on the paleontology of cerebral dominance

Observations on petalial asymmetry for 190 hominoid endocasts are reported, and their statistical differences assessed. While all taxa of hominoids show asymmetries to various degrees, the patterns or combinations of petalial asymmetries are very different, with fossil hominids and modern Homo sapiens showing an identical pattern of left-occipital, right-frontal petalias, which contrasts with those found normally in pongids. Of the pongids, Gorilla shows the greater degree of asymmetry in left-occipital petalias. Only modern Homo and hominids (Australopithecus, Homo erectus, Neandertals) show a distinct left-occipital, right-frontal petalial pattern. Analysis by x² statistics shows the differences to be highly significant. Due to small sample size and incompleteness of endocasts, small-brained hominids, i.e., Australopithecus, are problematical. To the degree that gross petalial patterns are correlated with cognitive task specialization, we speculate that human cognitive patterns evolved early in hominid evolution and were related to selection pressures operating on both symbolic and spatiovisual integration, and that these faculties are corroborated in the archaeological record.     

Fossil primates and the evolution of some primate locomotor systems

Of Paleocene primates only Plesiadapis is complete enough to reconstruct locomotor patterns; it was an arboreal scrambler, perhaps functioning like a large squirrel. Eocene lemurs (adapids) show an array of locomotor types much like certain modern Malagasy lemurs. The European Eocene tarsiid Necrolemur and the American Hemiacodon show the beginning of saltatory specializations in possession of elongated calcaneum and astragalus. Although not a direct anthropoid ancestor Necrolemur seems one of the best models for representing the early locomotor type from which higher primates arose. The Oligocene primates of Egypt (among which are the earliest undoubted pongids) are preserved with a forest fauna. Structures of long bones suggest they were arboreal. A considerable number of Miocene ape bones are known and those of Pliopithecus and Dryopithecus indicate similar adaptations. Of African Miocene forms, Dryopithecus major was a large, gorilla-sized animal, and hence perhaps primarily terrestrial. D. africanus was somewhat more arboreally adapted and a partial brachiator. The Italian fossil Oreopithecus, a coal-swamp dweller, shows indications of bipedality in pelvic structure. Ramapithecus, which is presumably ancestral to Australopithecus, shows palatal and facial patterns much like these later hominids, and probably hence had locomotor patterns more like men than like living apes; its lack of the dental specializations of apes strongly supports this suggestion.     

Humeral outlines in some hominoid primates and in Plio-Pleistocene hominids

A method of drawing outlines of the distal end of the humerus is presented and carried out on some pongids (Pan troglodytes, Pan paniscus, Pongo pygmaeus), on modern man, and on some casts of Plio-Pleistocene hominids. It appears that these outlines are good indicators of the overall morphology and permit the distinguishing of the different hominoids. For example, the morphology of the pillars surrounding the fossa olecrani is useful for this purpose. In modern man, the lateral pillar is quadrangular, contrasting with the triangular medial one. In pongids, both of them are triangular; however, it is possible to note differences between Pongo and Pan. In the South Asian ape, there is a stronger anteroposterior flattening of the pillars as well as the diaphysis. The similarity of the shape of the pillars might be considered as a result of an adaptation to suspension. The differences might be due to different weights of the animals. Plio-Pleistocene hominids are variable with regard to the morphology of this region. For example, Gombore IB 7594 is similar to Homo. KNM ER 739 exhibits features intermediate between hominids and pongids. Finally, AL 288.1M is closer to pongids. These results confirm a previous anatomical work.     

Functional implications of primate enamel thickness.

Recent evolutionary interpretations of Hominoidea have postulated functional relationships between tooth form, diet and masticatory biomechanics. A major consideration is the durability of the tooth under certain dietary conditions. Teeth with low cusps and thicker enamel are able to withstand heavy mastication of abrasive food bolus for a longer period. When comparisons are made between species of higher primates the variables of tooth size, cusp morphology, and enamel thickness appear to be related but until now no systematic analysis has been made to determine the functional relevance of several dental dimensions. This study provides data gained from comparisons of dentition of nine species of primates. Histological sections were made of the post canine teeth and 21 dimensions were compared. The relevant dimensions identified serve to withstand dental wear. The distribution of thicker enamel corresponded to the observed wear planes. Humans had thicker enamel than pongids while the macaque had the thinnest. These preliminary results tend to support theories which explain low, thick, enameled cusps in hominids.     

Affinity,hominid evolution and Creationism. New computer methods weaken the Creationists' position

Using samples of skulls of early and modern hominids and modern pongids we compare the plausibility of Evolutionist and Creationist viewpoints. Multivariate methods are used to assess the corollary of Creationism that all fossil specimens which Evolutionists consider to be hominids are either apes or humans. We conclude that the results of this study cannot easily be accommodated within a Creationist framework, and, therefore, contribute substantial evidence for the Evolutionist viewpoint.     

A Phylogenetic Study of Late Growth Events in a Mammalian Evolutionary Radiation—The Cranial Sutures of Terrestrial Artiodactyl Mammals

We recorded the relative timing of fusion of 29 ectocranial sutures in 480 skulls belonging to 35 extant and four fossil species from all major clades of terrestrial artiodactyls. The resulting data were studied in a phylogenetic context, using mapping of event-pairing of suture fusion events and Parsimov. As phylogenetic framework we generated a compound phylogeny from several previously published analyses. Overall suture closure per species ranged from five to all 29 sutures in Hexaprotodon. All living non-ruminants (suids, camelids, and hippopotamids) fuse more than 50% of the studied sutures (most over 75%), whereas in almost all ruminants less than 50% of the sutures fuse completely. Phylogenetic regression found a significant correlation between suture closure and body mass. In all species we observed an early fusion of the sutures surrounding the foramen magnum (Exoccipital-Supraoccipital, Exoccipital-Basioccipital), a consistent scheme also among other mammals. Scaling the number of changes to the number of sutures in each of the usually recognized skull modules reveals relatively equal numbers of changes in the cranial vault, the zygomatic-sphenoid region, the orbit, and the anterior oral-nasal region. Only the basicranium shows a much smaller number in terms of absolute and relative amount of suture fusion change. Some species show a unique pattern of suture fusion, such as the early fusion of many sutures in the palatal region in Pecari or that of premaxillary sutures in †Cainotherium, perhaps related to feeding mode. A strategy to strengthen the skull by obliterating the sutures could explain the pattern of increased sutural fusion in ruminant species with large cranial appendages.     

Comparative nutrient extraction from forages by grazing bovids and equids: a test of the nutritional model of equid/bovid competition and coexistence

Summary Ruminants are unevenly distributed across the range of body sizes observed in herbivorous mammals; among extant East African species they predominate, in numbers and species richness, in the medium body sizes (10–600 kg). The small and the large species are all hind-gut fermenters. Some medium-sized hind-gut fermenters, equid perissodactyls, coexist with the grazing ruminants, principally bovid artiodactyls, in grassland ecosystems. These patterns have been explained by two complementary models based on differences between the digestive physiology of ruminants and hind-gut fermenters. The Demment and Van Soest (1985) model accounts for the absence of ruminants among the small and large species, while the Bell/Janis/Foose model accounts both for the predominance of ruminants, and their co-existence with equids among the medium-sized species (Bell 1971; Janis 1976; Foose 1982). The latter model assumes that the rumen is competitively superior to the hind-gut system on medium quality forages, and that hind-gut fermenters persist because of their ability to eat more, and thus to extract more nutrients per day from high fibre, low quality forages. Data presented here demonstrate that compared to similarly sized grazing ruminants (bovids), hind-gut fermenters (equids) have higher rates of food intake which more than compensate for their lesser ability to digest plant material. As a consequence equids extract more nutrients per day than bovids not only from low quality foods, but from the whole range of forages eaten by animals of this size. Neither of the current nutritional models, nor refinements of them satisfactorily explain the preponderance of the bovids among medium-sized ungulates; alternative hypotheses are presented.     

Fractal dimensions of cranial sutures and waveforms.

Two quite different shapes of cranial sutures ostensibly yield fractal dimensions. The rare, intricate sutures yield the more valid fractal dimensions because self-similar scaling provides a double-log plot of negative slope. These sutures are fractals over a range of several r values. Some of the highly folded, wavy sutures in humans also fill space except at very tiny values of r, but are nonfractal. A great deal depends on whether the dimension D is > 1 and by how much, whether the curve yields a false fractal dimension, whether the curve scales and shows self-similarity, and whether the scaling occurs regularly in the same pattern. We suggest careful attention to the inverse power law equations, which when misused can yield false fractal values. Cranial sutures vary from the simple wavy sutures to the complex folded ones, and, in rare instances, evolve and develop to the self-similar, scaling, elaborate ones called intricate sutures. The main thing is to express the biology precisely, whether waveform regularity or irregularity or scaling elaboration conserving space and the original shape. D values may not in themselves reliably allow such a distinction, by whatever method used.     

Analysis of fractal dimension of O2A glial cells differentiating in vitro

Fractal dimension is a quantitative measure of morphological complexity. Glial cells of the oligodendrocyte-type 2 astrocyte (O2A) lineage exhibit increasing morphological complexity as they differentiate in vitro. Enriched populations of O2A progenitor cells isolated from neonatal rat cerebral hemispheres or optic nerves were allowed to differentiate in vitro, and their fractal dimensions were measured over time. The fractal dimensions of the maturing cells correlated with perceived complexity; cells with elaborate process branching had larger fractal dimensions than cells with a simpler morphology. An analysis of changes in fractal dimension revealed distinct rates of growth for both oligodendrocytes and type 2 astrocytes. The fractal dimension remained constant over a 10-fold range in optical magnification, demonstrating that cultured O2A glial cells exhibit self-similarity, a defining characteristic of fractal objects. These results illustrate that fractal dimension analysis of maturing cell populations is a useful method for quantitatively describing the process of cell differentiation.     

Basicranial flexion,relative brain size,and facial kyphosis in Homo sapiens and some fossil hominids

Comparative work among nonhominid primates has demonstrated that the basicranium becomes more flexed with increasing brain size relative to basicranial length and as the -upper and lower face become more ventrally deflected (Ross and Ravosa [1993] Am. J. Phys. Anthropol. 91:305–324). In order to determine whether modern humans and fossil hominids follow these trends, the cranial base angle (measure of basicranial flexion), angle of facial kyphosis, and angle of orbital axis orientation were measured from computed tomography (CT) scans of fossil hominids (Sts 5, MLD 37/38, OH9, Kabwe) and lateral radiographs of 99 extant humans. Brain size relative to basicranial length was calculated from measures of neurocranial volume and basicranial length taken from original skulls, radiographs, CT scans, and the literature. Results of bivariate correlation analyses revealed that among modern humans basicranial flexion and brain size/basicranial length are not significantly correlated, nor are the angles of orbital axis orientation and facial kyphosis. However, basicranial flexion and orbit orientation are significantly positively correlated among the humans sampled, as are basicranial flexion and the angle of facial kyphosis. Relative to the comparative sample from Ross and Ravosa (1993), all hominids have more flexed basicrania than other primates: Archaic Homo sapiens, Homo erectus, and Australopithecus africanus do not differ significantly from Modern Homo sapiens in their degree of basicranial flexion, although they differ widely in their relative brain size. Comparison of the hominid values with those predicted by the nonhominid reduced major-axis equations reveal that, for their brain size/basicranial length, Archaic and Modern Homo sapiens have less flexed basicrania than predicted. H. erectus and A. africanus have the degree of basicranial flexion predicted by the nonhominid reduced major-axis equation. Modern humans have more ventrally deflected orbits than all other primates and, for their degree of basicranial flexion, have more ventrally deflected orbits than predicted by the regression equations for hominoids. All hominoids have more ventrally deflected orbital axes relative to their palate orientation than other primates. It is argued that hominids do not strictly obey the trend for basicranial flexion to increase with increasing relative brain size because of constraints on the amount of flexion that do not allow it to decrease much below 90°. Therefore, if basicranial flexion is a mechanism for accommodating an expanding brain among non-hominid primates, other mechanisms must be at work among hominids. © 1995 Wiley-Liss, Inc.     

Analysis of a bone assemblage made by chimpanzees at Gombe National Park, Tanzania

Chimpanzee hunting provides information on prey characteristics and constraints acting on a large-bodied primate lacking a hunting technology, and has important implications for modeling hunting by fossil hominids. Analysis of the remains of five red colobus monkeys captured and consumed by Gombe chimpanzees in a single hunting bout provides one of the first opportunities to investigate the characteristics of prey bones surviving chimpanzee consumption. Four of the five individuals (an older infant, two juveniles and one subadult) were preserved in the bone assemblage; a neonate was entirely consumed. Cranial and mandibular fragments had the highest survivorships, followed by the scapulae and long bones. Post-cranial axial elements had the lowest survivorships. A high percentage (80%) of the long bones and ribs surviving consumption were damaged, most commonly through crenulation and step fracturing of bone ends. One of two partially reconstructed crania preserves a canine puncture through its left parietal. Proposed characteristics of faunal assemblages formed through chimpanzee-like hunting include small modal prey size, limited taxonomic diversity, a high proportion of immature individuals and a high frequency of skull bones. These characteristics would not uniquely identify hunting by fossil primates in the geological record, necessitating a contextual approach to diagnose hunting by hominids not forming an archeological record.Hominid utilization of vertebrate tissue is first unambiguously documented at 2.5 m.y.a. Rather than representing a strict "scavenging phase" in the evolution of hominid-prey interactions, Oldowan hominid carnivory may represent the overlay of large mammal scavenging on a tradition of small mammal hunting having a low archeological visibility.