A hominid tooth from Bulgaria: the last pre-human hominid of continental Europe

A hominid upper premolar was discovered in the Azmaka quarry, near Chirpan (Bulgaria). The associated fauna, especially the co-occurrence of Choerolophodon and Anancus among the proboscideans, and Cremohipparion matthewi and Hippotherium brachypus among the hipparions, constrains the age of the locality to the second half of the middle Turolian (ca. 7 Ma), making it the latest pre-human hominid of continental Europe and Asia Minor. The available morphological and metric data are more similar to those of Ouranopithecus from the Vallesian of Greece than to those of the early to middle Turolian hominids of Turkey and Georgia, but the time gap speaks against a direct phyletic link, and Turolian migration from the east cannot be rejected.     

Balance of the head in hominid evolution

In primates it is useful to distinguish three basic types of bipedal posture: (1) agonial, with extended hips and knees as in modern humans, (2) monogonial, with flexed hips but extended knees. and (3) digonial, with flexed hips and knees as in pongids. Early hominids retained an ancestral, forwardly inclined posture of the neck and head. Therefore the body posture of australopithednes must have differed from that in modem man, in which the centre of gravity of the head can be aligned with that of the body, other major centra of gravity, and important axes of rotation in a single frontal plane. It is suggested that in australopithednes the gravitational tilt of the head was counterbalanced by bent hips in association with hyperextended knees (monogonial posture). In australopithecines the increase in brain weight would have counteracted an improvement in the balance of the head. After the neck had assumed a more vertical posture as a consequence of shortening of the face, selection for an improved balance system in the bipedal posture favoured an increase in the weight of the postcondylar portion of the head, accentuated by selection for a posterior shift of the superior nuchal line in order to minimise the force of the nuchal muscles. At this stage the evolutionary increase in brain weight may have been largely a by-product of the process towards perfecting the bipedal posture. When the centre of gravity of the head had first become aligned with that of the body, the conditions of balance of the head had become favourable for a dramatic increase of brain size, as a result of selection for greater learning and storage capacity of the brain.     

Interproximal grooving in the Atapuerca-SH hominid dentitions

The dental sample recovered from the Sima de los Huesos (SH) Middle Pleistocene cave site of the Sierra de Atapuerca (Spain) includes 296 specimens. Interproximal wear grooves have been observed in 20 maxillary and mandibular posterior teeth belonging to at least five of the 32 individuals identified so far in the SH hypodigm. Interproximal grooving affected only the adults, and at an age between 25 and 40 years. The appearance, morphology, and location pattern of the SH wear grooves are similar to those reported in other fossil hominids and in more recent human populations. Two alternative proposals, the toothpicking and the fiber or sinew processing hypotheses, compete for explaining the formation of this anomalous wear. The characteristics observed in the wear grooves of the SH teeth are compatible only with the habitual probing of interdental spaces by means of hard and inflexible objects. Dietary grit may also have contributed to the abrasion of the root walls during the motion of the dental probes. Am. J. Phys. Anthropol. 102:369–376, 1997. © 1997 Wiley-Liss, Inc.     

Bone breakage in the Krapina hominid collection

The fragmentary condition of the Krapina Neandertal remains has been offered as one line of evidence for the hypothesis that these hominids were the victims of cannibals seeking marrow and brains. Two other hypotheses regarding the causes of the framentation have been raised: a substantial portion of the breakage in the Krapina collection is attributable to excavation damage; and the rest of the breakage is attributable to sedimentary pressure and to natural rock falls that occurred during the site's prehistory. The purpose of this paper is report on tests of these three hypotheses concerning the cause of breakage in the Krapina material. Microscopic inspection of all Krapina hominid specimens showed that 23% of the material was inadvertantly broken during excavation or during quarrying that took place at the end of the last century. The morphology of the prehistoric breakage is inconsistent with the cannibalism hypothesis and supports the hypothesis that prehistoric breakage was caused by sedimentary pressure and/or roof falls.     

Cranial capacity in hominid evolution

We present an analysis of cranial capacity of 118 hominid crania available from the literature. The crania belong to both the genusAustralopithecus andHomo and provide a clear outline of hominid cranial evolution starting at more than 3 million years ago. Beginning withA. afarensis there is a clear increase in both absolute and relative brain size with every successive time period.H.s. neandertal has an absolutely and relatively smaller brain size (1412cc, E.Q.=5.6) than fossil modernH.s. sapiens (1487cc, E.Q.=5.9). Three evolutionary models of hominid brain evolution were tested: gradualism, punctuated equilibrium, and a mixed model using both gradualism and punctuated equilibrium. Both parametric and non-parametric analyses show a clear trend toward increasing brain size withH. erectus and a possible relationship within archaicH. sapiens. An evolutionary stasis in cranial capacity could not be refuted for all other taxa. Consequently, the mixed model appears to more fully explain hominid cranial capacity evolution. However, taxonomic decisions could directly compromise the possibility of testing the evolutionary mechanisms hypothesized to be operating in hominid brain expansion.     

Variability selection in hominid evolution

Variability selection (abbreviated as VS) is a process considered to link adaptive change to large degrees of environment variability. Its application to hominid evolution is based, in part, on the pronounced rise in environmental remodeling that took place over the past several million years. The VS hypothesis differs from prior views of hominid evolution, which stress the consistent selective effects associated with specific habitats or directional trends (e.g., woodland, savanna expansion, cooling). According to the VS hypothesis, wide fluctuations over time created a growing disparity in adaptive conditions. Inconsistency in selection eventually caused habitat-specific adaptations to be replaced by structures and behaviors responsive to complex environmental change. Key hominid adaptations, in fact, emerged during times of heightened variability. Early bipedality, encephalized brains, and complex human sociality appear to signify a sequence of VS adaptations—i.e., a ratcheting up of versatility and responsiveness to novel environments experienced over the past 6 million years. The adaptive results of VS cannot be extrapolated from selection within a single environmental shift or relatively stable habitat. If some complex traits indeed require disparities in adaptive setting (and relative fitness) in order to evolve, the VS idea counters the prevailing view that adaptive change necessitates long-term, directional consistency in selection. © 1998 Wiley-Liss, Inc.     

Genetic aspects in hominid evolution

Genomic comparison between apes and humans have made important contributions to our understanding of human evolution. The modern period of karyological comparisons between humans and other primates began about forty years ago and has been marked by a series of technical revolutions. In the 1960s pioneering genetic and chromosomal comparisons of human and great apes suggested, as had Darwin a century before, that our closest relative were the African apes. Early immunological analyses placed human/apes divergence at about five million year ago. Acceptance of man’s late divergence from the African apes was delayed by the scarcity of paleontological evidence coupled with a fallacious Asiatic origin hypothesis of the hominoids. Chromosome banding techniques in the seventies and high resolution methods in the eighties allowed a detailed comparison of the chromosomes between closely related primates and reinforced the hypothesis of an African origin for humans. It was clearly shown that humans were more closely related to African apes than to the orang-utan. The last decade has seen a vigorous integration of molecular and cytogenetic. This powerful combination promises to be quite fruitful because chromosomes can be compared directly at the DNA level. Fluorescentin situ hybridisation (FISH), chromosome painting, is a colourful technique for establishing chromosomal homology between species. Results obtained by FISH over the last ten years have resolved the cytogenetic problem of the homology between humans, apes, hylobates and Old World monkeys and defined the chromosomal syntenies and major translocations involved in the genome evolution of higher primates.     

Bioenergetics and the origin of hominid bipedalism

Compared to most quadrupedal mammals, humans are energetically inefficient when running at high speeds. This fact can be taken to mean that human bipedalism evolved for reasons other than to reduce relative energy cost during locomotion. Recalculation of the energy expended during human walking at normal speeds shows that (1) human bipedalism is at least as efficient as typical mammalian quadrupedalism and (2) human gait is much more efficient than bipedal or quadrupedal locomotion in the chimpanzee. We conclude that bipedalism bestowed an energetic advantage on the Miocene hominoid ancestors of the Hominidae.     

Palaeoenvironments and the origin of hominid bipedalism

Abstract

It has long been accepted that hominids emerged during the Pliocene in a savannah environment in which a terrestrial quadruped gradually developed bipedal adaptations. However, data from the Late Miocene (i.e. 7–7.5 Ma), including detailed palaeontological and biogeochemical studies, suggest that our earliest Upper Miocene ancestors inhabited well-wooded to forested environments where they could have spent a certain amount of time in the trees. A plausible type of ecosystem in which upright posture and bipedal locomotion could have emerged is represented by Miombo Woodland, in which vertical arboreal supports predominate and trees are separated from each other by gaps. Subsequently hominids dispersed into the Savannah as accomplished bipeds, but retained the ability to climb trees. This scenario is compatible with the postcranial anatomy of Australopithecus, including its femoral elongation, body proportions, manual precision grip (also present in 6-million-year-old Orrorin) and a non-prehensile hallux.     

Cladistics and the hominid fossil record

Cladistic methodology has become common in phylogenetic analyses of the hominid fossil record. Even though it has correctly placed emphasis on morphology for the primary determination of affinities between groups and on explicit statements regarding traits and methods employed in making phylogenetic assessments, cladistics nonetheless has limitations when applied to the hominid fossil record. These include 1) the uncritical assumption of parsimony, 2) uncertainties in the identification of homoplasies, 3) difficulties in the appropriate delimitation of samples for analysis, 4) failure to account for normal patterns of variation, 5) methodological problems with the appropriate identification of morphological traits involving issues of biological relevance, intercorrelation, primary versus secondary characters, and the use of continuous variables, 6) issues of polarity identification, and 7) problems in hypothesis testing. While cladistics has focused attention on alternative phylogenetic reconstructions in hominid paleontology and on explicit statements regarding their morphological and methodological underpinnings, its biological limitations are too abundant for it to be more than a heuristic device for the preliminary ordering of complex human paleontological and neonatological data.     

Multiple dispersals and the environment in hominid evolution

Conclusions Given that we have attempted a very broad sweep through two of the major issues in anthropology (human evolution and environmental change) it would be impossible to have any easy conclusions. In closing I would therefore stress only the major points in their simplest form. First, hominid evolution does not consist of just two dispersals (out of Africa 1 and 2) but of multiple such events. Unraveling these, and the ecological conditions underlying them, should be a major focus for research. Second, if hominid evolution does consist of multiple dispersals, then it follows that there must have been multiple local extinctions. Although we tend to think of human demography in terms of growth, demographic collapse is equally important in evolution. Third, where hominids have had an effect on the environment it has tended to occur in two very specific contexts-the colonisation of new regions and the development of agriculture. And finally, if we are interested in the human evolution and the environment, coevolution rather than anthropogenic change may be the more useful concept.     

Affinities of the Swartkrans 847 hominid cranium

Further evidence of the presence of a second hominid species at the Swartkrans locality was obtained in 1969 when the SK.847 specimen was discovered by us to represent the same individual as the SK.80 maxilla. The SK.847 specimen had previously been regarded as robust australopithecine, whereas the latter was first attributed to Telanthropus capensis and subsequently to a species of the genus Homo. Recent criticism of our interpretation of these remains has not evaluated and analyzed critically the primary fossil evidence. Instead it relies on a strict adherence to an as yet unsubstantiated hypothesis that posits only a single hominid species at any point in space and time in the Cenozoic history of Hominidae.     

Biomechanical interpretation of the early hominid hip

A new pelvic fragment from Swartkrans provides the opportunity to analyze the hip joint mechanics of the robust form of early hominid. The function of the lateral support system provided by the abductor muscles of the hip appears to be similar to that of the gracile early hominid from Sterkfontein. The system is well adapted for providing the lateral support necessary for efficient bipedalism. The hip extensor mechanism and hip internal rotatory system also appear to be well adapted for efficient bipedalism in a way very similar to the other early hominids. The conclusion reached is that the robust and gracile forms of South African early hominids were basically similar in their locomotor adaptation and were most likely habitual bipeds.     

Ecological dominance and the final sprint in hominid evolution

In contrast to many other models of human evolution the “balance of power” theory of Alexander has a clear answer to the question why a runaway selection process for unique social and moral capacities occurred in our ancestry only and not in other species: “ecological dominance” is hypothesized to have diminished the effects of “extrinsic” forces of natural selection such that withinspecies, intergroup competition increased (Alexander, 1989). Alexander seems to be wrong, however, in his claim that already the common HUCHIBO (Humans, Chimps, Bonobo's)-ancestor has crossed the ecological dominance barrier. In this paper an adapted version of Alexander's model is presented and several different ways are proposed to make this adapted version testable. A preliminary survey of the available paleontological and paleoecological data suggests that there is some evidence of a less vulnerable position towards predators in earlyHomo and that there are clear signs related to a crossing of the ecological dominance barrier inHomo sapiens sapiens.     

Sociobiology of the great apes and the hominid ancestor

This review of recent field studies of the great apes summarizes and weighs socioecological and sociobiological evidence concerning the ultimate causes of social structure. The behavioral ecology and social structures of mountain gorillas, orangutans, chimpanzees, and bonobos are reviewed and contrasted between species. Social dynamics and molecular studies indicate that, among the extant Hominoidea, the evolutionary clade of chimpanzees, bonobos, and humans probably evolved from the most recent common ancestor in the ape-human stem. The most probable phylogenetic referential model for the suite of social behaviors of the hominid ancestor consists of the behavioral traits common to all three species: female exogamy, male retention, female associations due to attraction to the same male(s), weak bonds between females, a closed, stable social group made up of a kin-group of males and containing multiple females, fusion-fission sociality in which individuals of either sex sometimes travel alone, a polygynous mating system, communal territoriality with cooperative defense by kin-related males who exhibit strong solidarity among themselves but who may kill other males in territorial disputes, low mating competition between males within communities, and moderate sexual dimorphism. It is postulated that this phylogenetic model is a useful tool for comparing goodness of fit of other referential models seeking to explain hominid evolution. It is also suggested that to construct a “strategic” or conceptual model to explain hominid evolution, the putative evolutionary processes responsible for this male-retentive system require further testing in the field by measuring individual reproductive success among the great apes and man.