Observed and expected variation in hominid evolution

All of the major groups of fossil hominids (australopithecines, pithecanthropines, Neandertals, and early sapiens) were discovered by 1925, and therefore prior to the formulation of the synthetic theory of evolution that revolutionized the concept of the species in systematics. While these fossil finds were being made the framework for their interpretation included several assumptions: (1) that the number of living hominoid species was great, and that intraspecific variation was slight (authoritative sources recognized as many as 14 separate species of chimpanzees and 15 species of gorillas); (2) that the timescale of human evolution was brief (measured in tens or hundreds of thousands of years). As a result of these premises the consensus that hominid evolution was characterized by a large number of sympatric and synchronic species was virtually inevitable.In contrast, recent molecular studies demonstrate that genetic diversity among recent hominoids is so slight that even humans and chimpanzees differ at only about 1% of the loci that have been sampled so far; evidently, very small genetic differences can produce rather great contrasts in morphology. At the same time, geological break-throughs have increased the timescale for human evolution to several million years.It is concluded that morphological differences among fossil hominids, even if very appreciable and complex, do not necessarily reflect a great degree of either genetic or taxonomic diversity. Potential effects of evolutionary change through time should be incorporated into models of hominid evolution as a means of assessing the minimum number of lineages required to account for observed variations among hominid specimens.     

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.     

Can revisionism,in evolutionary biology,help in formulating hypotheses about hominid evolution?

While evolutionary theory has been undergoing considerable revision for more than a decade scarcely any biological scientists are willing to resurrect the notion of acquired inheritance since it totally jndermines the concept of ancestor-descendant stability. Acquired inheritance is divisible into two components, soma to germ transfer to replicable information and directed or anticipatory mutation. Pure soma to germ transfer still allows for random events or natural selection as is evident from E. Steel’s proposal of the “somatic selection hypothesis”. Directed mutation is non-stochastic and self-referent. Recent bacterial research has claimed strong support for directed mutation but this phenomenon still would not be important for the small proportion of living taxa (including primates) who sequester their germlines early in ontogeny. There is a basic incompatibility between purposeful responses to final growth stages in complex metazoans and the need to permanently fix future growth into heritable programs. Symposium on Foundations for different approaches to the study of human evolution September 1–3, 1989, Liblice, Czechoslovakia, Czechoslovak Academy of Science     

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.     

The genetic implications of biological and cultural structuring processes during hominid evolution

Human populations are genetically structured through biological reproduction but reproduction is socially structured through culturally expressed systems of marriage, hence the need to take into account the cultural dimension of human societies when considering the genetic structure of modern human populations. The interplay between these two structuring processes is examined through the implications of a biologically based primate form of social organization versus a culturally based foraging form of social organization for an anomaly between the genetic differentiation implied by micro-evolutionary genetic models versus the species-wide pattern of morphological change implied by hominid fossil evidence. Simulation is used to derive the pattern for genetic differentiation at the level of the living group for both forms of social organization and a competition model is added to the genetic model as a way to resolve the anomaly. The importance of the competition model for the transition from a biologically based to a culturally based form of social organization is discussed.     

Analysis of an early hominid ulna from the Omo basin,Ethiopia

The discovery (in 1971) of a nearly complete right ulna from the Shungura Formation of the Omo basin provides the opportunity to analyze the forelimb structure of the Australopithecus boisei form of early hominid. Results from multivariate morphometric analyses show that this bone is unique in shape among the extant hominoids although it is most similar to Pan and Homo. Despite its long slender shaft and large distal articular surface the bone's overall morphology is quite unlike Pongo.     

The environmental background of hominid emergence and the appearance of the genusHomo

The human biologist usually considers ecology of recent humanity. This essay explores the question of whether the human biologist specialising in the ecology of living peoples has anything to learn from the palaeo-anthropologist, studying ancient hominids andtheir adaptive mechanisms over a deep time dimension. Since the Hominidae are under discussion, the definition of the hominids is reviewed. Historically, three phases are recognised. A rethinking of the classification of the hominoids has become necessary for the old and classical systematics, which divided this superfamily into the Hominidae and the Pongidae, is now outmoded. Since no consensus on such a re-classification has yet been reached, the author adheres to the classical system for the time being. The Hominidae emerged between about 8 and 5 million years ago. At that time, Africa was subject to major cooling and aridification and considerable changes in the flora and fauna were occurring. Wet forests were retreating, savanna was spreading and the animals of Africa were undergoing many changes, partly by faunal interchange with Asia following the drying up of the Mediterranean, and partly by autochthonous evolution among the pre-existing species of the continent. The Hominidae could well have emerged from the striking environmental modifications of this late Miocene phase. Critical changes occurred in hominid evolution between 3 and 2 million years before the present. The pre-existing speciesAustralopithecus africanus acquired the form of a postulated derivedA. africanus; the hominid lineage underwent cladogenetic splitting into robust and hyper-robust australopithecines and the genusHomo; Homo babilis appeared; stone tools are first found in the archaeological record; spoken language seems to have been acquired. These sensational events, within the space of one million years, took place against the background of conspicuous changes in the climate and physical geography of Africa, the flora and non-hominid fauna. Mankind became increasingly dependent upon stone culture. Hence a new element was added to the range of modes of adjustment, an element which must have greatly increased the ecological flexibility of the hominids. From the end of the Pliocene era onwards, culture should be seen as a constituent of man's environment and, at the same time, a highly advantageous component of human adaptational processes. In later and recent mankind, it may be difficult to extricate the respective roles of biological, social and physical factors, on the one hand, and cultural aspects on the other, as mechanisms and facilitators of adaptation to diverse econiches.     

An integrated approach to taphonomy and faunal change in the Shungura formation (Ethiopia) and its implication for hominid evolution

Environmental and faunal changes through time have been recorded for many African Plio-Pleistocene sites. Fossil evidence suggests that there is a continuous, if not uniform, transformation of the fauna and flora from the Pliocene through the end of Pleistocene. However, discerning major biotic turnovers and linking them to global and regional climatic changes have been complicated by many factors, notably taphonomy and discontinuity of the fossil evidence, notwithstanding the considerable work of some researchers (e.g., Vrba, E.S., 1988. Late Pliocene climatic events and hominid evolution, in: Grine, F. (Ed.), Evolutionary History of the "Robust" Australopithecines. De Gruyter, New York, pp. 405-426, Vrba, E.S., 1995. The fossil record of African (Mammalia, Bovidae) in relation to human evolution and paleoclimate, in: Vrba, E.S., Denton, G.H., Partridge, T.C., Burkle, L.H. (Eds.), Paleoclimate and Evolution, with Emphasis on Human Origins. Yale University Press, New Haven, pp. 385-424). A sample of over 22,000 fossils collected by the French Omo Expedition, from the Shungura Formation of Ethiopia, was analyzed using an integrated approach to investigate taphonomic and faunal change patterns. The following results are obtained: (1) Univariate and multivariate studies support continuous faunal change from Member A through Member G of the Shungura sequence; (2) Correspondence analysis (CA) on extant bovids in African game parks shows that bovid tribes and genera are generally characterized by habitat specificity; (3) Taphonomic studies demonstrate that the relative abundance of different skeletal elements varies according to depositional environment; (4) CA on 73 localities of the Shungura Formation and 19 mammalian taxa points to a major faunal change around the base of Member G dated to ca. 2.3 Ma. This transformation is characterized by a change to open and edaphic grassland as a dominant type of environment; (5) This major faunal change correlates in time with the appearance of A. boisei. It is tentatively suggested that this major biome change is associated with an anagenetic speciation from A. aethiopicus to A. boisei.     

Comparison of inverse-dynamics musculo-skeletal models of AL 288-1 Australopithecus afarensis and KNM-WT 15000 Homo ergaster to modern humans, with implications for the evolution of bipedalism

Size and proportions of the postcranial skeleton differ markedly between Australopithecus afarensis and Homo ergaster, and between the latter and modern Homo sapiens. This study uses computer simulations of gait in models derived from the best-known skeletons of these species (AL 288-1, Australopithecus afarensis, 3.18 million year ago) and KNM-WT 15000 (Homo ergaster, 1.5-1.8 million year ago) compared to models of adult human males and females, to estimate the required muscle power during bipedal walking, and to compare this with those in modern humans. Skeletal measurements were carried out on a cast of KNM-WT 15000, but for AL 288-1 were taken from the literature. Muscle attachments were applied to the models based on their position relative to the bone in modern humans. Joint motions and moments from experiments on human walking were input into the models to calculate muscle stress and power. The models were tested in erect walking and 'bent-hip bent-knee' gait. Calculated muscle forces were verified against EMG activity phases from experimental data, with reference to reasonable activation/force delays. Calculated muscle powers are reasonably comparable to experimentally derived metabolic values from the literature, given likely values for muscle efficiency. The results show that: 1) if evaluated by the power expenditure per unit of mass (W/kg) in walking, AL 288-1 and KNM-WT 15000 would need similar power to modern humans; however, 2) with distance-specific parameters as the criteria, AL 288-1 would require to expend relatively more muscle power (W/kg.m(-1)) in comparison to modern humans. The results imply that in the evolution of bipedalism, body proportions, for example those of KNM-WT 15000, may have evolved to obtain an effective application of muscle power to bipedal walking over a long distance, or at high speed.     

The mixed dentition and associated skull fragments of a juvenile fossil hominid from Sterkfontein,South Africa

In April–May 1983, the late A.R. Hughes and his field team recovered more than 40 bone fragments and teeth from a single solution pocket of the Sterkfontein Formation. After preparation and reconstruction by JMC, it was recognised that these fragments represent a single juvenile individual (Stw 151), consisting of more than 40 cranial and dental parts, with mixed dentition. It constitutes the most complete set of jaws and teeth of an early hominid child since the Taung child was recovered in 1924. In this paper, the morphological and metrical features of the individual teeth are described. The other associated skull fragments (right ramus of the mandible, left petrous bone, right glenoid region) are also described. Comparisons are made with other South (and East) African fossil hominids. The beautiful preservation simultaneously of most of the deciduous teeth and of the permanent teeth exposed in their crypts allows an accurate analysis of the developmental sequence. A report on the dental developmental status of this juvenile is presented. On the basis of the microanatomical study of the developing permanent teeth, the estimated age at death is 5.2–5.3 years. Reconstructions of the maxillary and mandibular arcades are also offered. The morphological and metrical features of Stw 151 raise the possibility that it may represent a hominid more derived towards an early Homo condition than the rest of the A. africanus sample from Member 4. Am J Phys Anthropol 106:425–465, 1998. © 1998 Wiley-Liss, Inc.     

New Late-Pleistocene uranium–thorium and ESR dates for the Singa hominid (Sudan)

The Singa (Sudan) calvaria has been interpreted previously as a terminal Pleistocene modern human fossil, perhaps related to the Bushman of Southern Africa. Here we report new mass-spectrometric U–Th dates for the calcrete deposit enclosing the fossil teeth and the calvaria itself and new electron spin resonance (ESR) dates for associated dental materials. The new data constrain the age of the hominid to at least 133±2 ka. Together with the preferred linear uptake (LU) ESR dates, the U–Th data confirm that the intriguing mixture of modern and archaic characteristics in the Singa specimen date from isotope stage 6. Far from being a modern human fossil, it represents a rare example of an archaic African population which may have been ancestral to all modernHomo sapiens.     

Science, religion, and society: the problem of evolution in america

American resistance to accepting evolution is uniquely high among First World countries. This is due largely to the extreme religiosity of the United States, which is much higher than that of comparably advanced nations, and to the resistance of many religious people to the facts and supposed implications of evolution. The prevalence of religious belief in the United States suggests that outreach by scientists alone will not have a huge effect in increasing the acceptance of evolution, nor will the strategy of trying to convince the faithful that evolution is compatible with their religion. Because creationism is a symptom of religion, another strategy to promote evolution involves loosening the grip of faith on America. This is easier said than done, for recent sociological surveys show that religion is highly correlated with the dysfunctionality of a society, and various measures of societal health show that the United States is one of the most socially dysfunctional First World countries. Widespread acceptance of evolution in America, then, may have to await profound social change.     

New discoveries and interpretations of hominid fossils and artifacts from Vindija Cave, Croatia

Beginning with excavations during the 1970s, Vindija Cave (Croatia) has yielded significant Middle and Upper Paleolithic fossil and archaeological finds. We report on seven recently identified hominid fossils, a newly associated partial hominid cranial vault from level G(3), nine possible bone retouchers, and a revised interpretation of the Mousterian artifact assemblage from the site. This new information reinforces our knowledge of the complex biocultural phenomena revealed in unit G and earlier deposits at Vindija. Six of the new hominid fossils derive from stratigraphic units G and I, while one lacks exact provenience. All specimens preserving diagnostic anatomy are from Neandertals. One of the postcranial remains, a radius fragment which exhibits Neandertal-like anatomy, comes from level G(1)and is congruent with the previously established association of Neandertals with an early Upper Paleolithic industry at the site. The partial cranial vault represents the most complete Neandertal from Vindija. The possible retouchers derive from unit G. Our analysis of these artifacts suggests that both percussion and pressure techniques may have been used by Neandertals in the final stage of tool production (retouching). This paper also presents a revision of the artifact analysis for late Mousterian level G(3). We separated raw materials into two main groups due to the differing ways that the materials fracture and the differing morphology of the debitage. The use of raw material in level G(3)is different from earlier Middle Paleolithic levels at Vindija. This indicates that the G(3)late Neandertals were making choices regarding source material somewhat more like the Upper Paleolithic people at the site. When interpreted within a larger regional framework, the Vindija archaeological and hominid fossil remains demonstrate a complex, mosaic pattern of biocultural change in the Late Pleistocene of south-central Europe.