Multiple dispersals and modern human origins

Despite a massive endeavour, the problem of modern human origins not only remains unresolved, but is usually reduced to “Out of Africa” versus multiregional evolution. Not all would agree, but evidence for a single recent origin is accumulating. Here, we want to go beyond this debate and explore within the “Out of Africa” framework an issue that has not been fully addressed: the mechanism by which modern human diversity has developed. We believe there is no clear rubicon of modern Homo sapiens, and that multiple dispersals occurred from a morphologically variable population in Africa. Pre-existing African diversity is thus crucial to the way human diversity developed outside Africa. The pattern of diversity—behavioural, linguistic, morphological and genetic—can be interpreted as the result of dispersals, colonisation, differentiation and subsequent dispersals overlaid on former population ranges. The first dispersals would have originated in Africa from where two different geographical routes were possible, one through Ethiopia/Arabia towards South Asia, and one through North Africa/Middle East towards Eurasia.     

From apes to humans: locomotion as a key feature for phylogeny

If bipedalism has often been considered to be of a crucial interest for understanding hominid evolution, the acceptance of locomotor features to build phylogenies is still far from being a reality in the field. Especially for hominid evolution, it still seems to be difficult to accept that traits, other than craniodental ones, can be useful for defining the major dichotomies. The recent discovery of Australopithecus anamensis suggests a challenging view of the major dichotomy between apes and humans. Whilst it is widely accepted that Ardipithecus ramidus is ancestral to Australopithecus anamensis, which in its turn is ancestral to Australopithecus afarensis and then to later hominids, the postcranial adaptations, which should be taken into account, suggest another branching pattern. Based on the fact that by 4.0 million years two different locomotor patterns can be identified in hominids, two lineages would appear to be present: the "Australopithecine" lineage (with Australopithecus afarensis or Ardipithecus ramidus if the latter is really a hominid sensu stricto) and the "Hominine" lineage (with Australopithecus anamensis = Praeanthropus africanus).     

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.     

Survival of the fattest: the key to human brain evolution

The circumstances of human brain evolution are of central importance to accounting for human origins, yet are still poorly understood. Human evolution is usually portrayed as having occurred in a hot, dry climate in East Africa where the earliest human ancestors became bipedal and evolved tool-making skills and language while struggling to survive in a wooded or savannah environment. At least three points need to be recognised when constructing concepts of human brain evolution : (1) The human brain cannot develop normally without a reliable supply of several nutrients, notably docosahexaenoic acid, iodine and iron. (2) At term, the human fetus has about 13 % of body weight as fat, a key form of energy insurance supporting brain development that is not found in other primates. (3) The genome of humans and chimpanzees is <1 % different, so if they both evolved in essentially the same habitat, how did the human brain become so much larger, and how was its present-day nutritional vulnerability circumvented during 5-6 million years of hominid evolution ? The abundant presence of fish bones and shellfish remains in many African hominid fossil sites dating to 2 million years ago implies human ancestors commonly inhabited the shores, but this point is usually overlooked in conceptualizing how the human brain evolved. Shellfish, fish and shore-based animals and plants are the richest dietary sources of the key nutrients needed by the brain. Whether on the shores of lakes, marshes, rivers or the sea, the consumption of most shore-based foods requires no specialized skills or tools. The presence of key brain nutrients and a rich energy supply in shore-based foods would have provided the essential metabolic and nutritional support needed to gradually expand the hominid brain. Abundant availability of these foods also provided the time needed to develop and refine proto-human attributes that subsequently formed the basis of language, culture, tool making and hunting. The presence of body fat in human babies appears to be the product of a long period of sedentary, shore-based existence by the line of hominids destined to become humans, and became the unique solution to insuring a back-up fuel supply for the expanding hominid brain. Hence, survival of the fattest (babies) was the key to human brain evolution.     

Divergent patterns of integration and reduced constraint in the human hip and the origins of bipedalism

When compared to other hominids--great apes including humans--the human pelvis reveals a fundamental reorganization of bony morphology comprised of multiple trait-level changes, many of which are associated with bipedal locomotion. Establishing how patterns of integration--correlations and covariances among traits--within the pelvis have evolved in concert with morphology is essential to explaining this evolutionary transition because integration may facilitate or constrain morphological evolution. Here, we show that the human hip bone has significantly lower levels of integration and constraint overall when compared to other hominids, that the focus of these changes is on traits hypothesized to play major functional roles in bipedalism, and we provide evidence that the human hip was reintegrated in a pattern distinct from other members of this group. Additionally, the evolutionary transition from a nonhuman great ape-like to human hip bone morphology was significantly easier to traverse using the human integration pattern in each comparison, which suggests hominin patterns may have evolved to facilitate this transition. Our results suggest natural selection for bipedalism broke down earlier hominid integration patterns, allowing relevant traits to respond to separate selection pressures to a greater extent than was previously possible, and reintegrated traits in a way that could have facilitated evolution along the vector specifying ancestral hominid and hominin morphological differences.     

Increased breathing control: Another factor in the evolution of human language

Investigation into the evolution of human language has involved evidence of many different kinds and approaches from many different disciplines. For full modern language, humans must have evolved a range of physical abilities for the production of our complex speech sounds, as well as sophisticated cognitive abilities. Human speech involves free‐flowing, intricately varied, rapid sound sequences suitable for the fast transfer of complex, highly flexible communication. Some aspects of human speech, such as our ability to manipulate the vocal tract to produce a wide range of different types of sounds that form vowels and consonants, have attracted considerable attention from those interested in the evolution of language. 1 , 2 However, one very important contributory skill, the human ability to attain very fine control of breathing during speech, has been neglected. Here we present evidence of the importance of breathing control to human speech, as well as evidence that our capabilities greatly exceed those of nonhuman primates. Human speech breathing demands fine neurological control of the respiratory muscles, integrated with cognitive processes and other factors. Evidence from comparison of the vertebral canals of fossil hominids and those of extant primates suggests that a major increase in thoracic innervation evolved in later hominid evolution, providing enhanced breathing control. If that is so, then earlier hominids would have had quite restricted speech patterns, whereas more recent hominids, with human‐like breath control abilities, would have been capable of faster, more varied speech sequences.     

The evolution and functions of laughter and humor: a synthetic approach

A number of recent hypotheses have attempted to explain the ultimate evolutionary origins of laughter and humor. However most of these have lacked breadth in their evolutionary frameworks while neglecting the empirical existence of two distinct types of laughter--Duchenne and non-Duchenne--and the implications of this distinction for the evolution of laughter as a signal. Most of these hypotheses have also been proposed in relative isolation of each other and remain disjointed from the relevant empirical literature. Here we attempt to remedy these shortcomings through a synthesis of previous laughter and humor research followed by (i) a reevaluation of this research in light of theory and data from several relevant disciplines, and (ii) the proposal of a synthetic evolutionary framework that takes into account phylogeny and history as well as proximate mechanisms and adaptive significance. We consider laughter to have been a preadaptation that was gradually elaborated and co-opted through both biological and cultural evolution. We hypothesize that Duchenne laughter became fully ritualized in early hominids between 4 and 2 mya as a medium for playful emotional contagion. This mechanism would have coupled the emotions of small hominid groups and promoted resource-building social play during the fleeting periods of safety and satiation that characterized early bipedal life. We further postulate that a generalized class of nonserious social incongruity would have been a reliable indicator of such safe times and thereby came to be a potent distal elicitor of laughter and playful emotion. This class of stimuli had its origins in primate social play and was the foundation for formal human humor. Within this framework, Duchenne laughter and protohumor were well established in the hominid biobehavioral repertoire when more cognitively sophisticated traits evolved in the hominid line between 2 mya and the present. The prior existence of laughter and humor allowed them to be co-opted for numerous novel functions, and it is from this process that non-Duchenne laughter and the "dark side" of laughter emerged. This perspective organizes the diversified forms and functions that characterize laughter and humor today and clarifies when and how laughter and humor evolved during the course of human evolution.     

Brief communication: Reaction to fire by savanna chimpanzees (Pan troglodytes verus) at Fongoli,Senegal: Conceptualization of “fire behavior” and the case for a chimpanzee model

The use and control of fire are uniquely human traits thought to have come about fairly late in the evolution of our lineage, and they are hypothesized to correlate with an increase in intellectual complexity. Given the relatively sophisticated cognitive abilities yet small brain size of living apes compared to humans and even early hominins, observations of wild chimpanzees' reactions to naturally occurring fire can help inform hypotheses about the likely responses of early hominins to fire. We use data on the behavior of savanna chimpanzees (Pan troglodytes verus) at Fongoli, Senegal during two encounters with wildfires to illuminate the similarities between great apes and humans regarding their reaction to fire. Chimpanzees' close relatedness to our lineage makes them phylogenetically relevant to the study of hominid evolution, and the open, hot and dry environment at Fongoli, similar to the savanna mosaic thought to characterize much of hominid evolution, makes these apes ecologically important as a living primate model as well. Chimpanzees at Fongoli calmly monitor wildfires and change their behavior in anticipation of the fire's movement. The ability to conceptualize the “behavior” of fire may be a synapomorphic trait characterizing the human‐chimpanzee clade. If the cognitive underpinnings of fire conceptualization are a primitive hominid trait, hypotheses concerning the origins of the control and use of fire may need revision. We argue that our findings exemplify the importance of using living chimpanzees as models for better understanding human evolution despite recently published suggestions to the contrary. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Ontogeny and phylogeny of femoro-tibial characters in humans and hominid fossils: functional influence and genetic determinism.

Three different human femoro-tibial characters are selected as functionally relevant and derived hominid characters: femoral bicondylar angle, shape of the femoral distal epiphysis, and the tibial insertion of the lateral meniscus. The timing and mode of formation of these characters are investigated during human ontogeny and are shown to differ considerably. The available hominid fossils (Australopithecus afarensis and early Homo) are interpreted in the light of this ontogenetic analysis with the conclusion that, during hominid evolution, different modes of selection of these features must have occurred. In modern humans, the femoral bicondylar angle proves to be an epigenetic functional feature, which develops during early childhood growth. It is present in all australopithecines and we suggest that it developed following a change in their locomotor behavior and not upon a genomic change: the early practice of bipedal walking, with adducted knee joints, in the locomotor repertoire of infant australopithecines, was sufficient to promote this angle. Later in hominid evolution, the knee joint evolved from having a single insertion of the lateral meniscus on the tibia to a double one. While Australopithecus afarensis exhibits a single insertion, early Homo clearly exhibits a double insertion of the lateral meniscus on the tibia. The double insertion restricts the mobility of the meniscus on the tibial plateau, indicating a habitual practice of full extension movements of the knee joint. Among modern humans, the posterior insertion of the lateral meniscus appears early in fetal life. Consequently in early Homo, this new selected feature developed directly as a result of a genomic change. The derived shape of human distal femoral epiphysis includes a prominence of the lateral lip of the femoral trochlea, an elliptical profile of the lateral condyle, and an anteroposterior lengthening of the epiphysis. Analysis of human fetal and neonatal distal epiphyses shows that the prominence of the lateral lip of the trochlea arises before any use, and thus appears to be genetically determined. However, the postnatal development of this joint shows that this feature is also modified epigenetically by use. It is argued that the hominid femoro-patellar joint would have been reshaped following the process of genetic assimilation (Waddington [1942] Nature 3811:563-565). The prominence of the lateral lip of the femoral trochlea was probably selected following a two-staged process-first epigenetic, then genetic. Far from being a Lamarckian explanation, this concept applies precisely to adaptive characters that are induced by an external stimulus during a single lifetime and are replaced through natural selection by genetically based equivalent characters. The nature of the structures involved in the studied features is shown to be an important parameter determining their mode of development and selection.     

The relative cost of bent-hip bent-knee walking is reduced in water

The debate about how early hominids walked may be characterised as two competing hypotheses: They moved with a fully upright (FU) gait, like modern humans, or with a bent-hip, bent-knee (BK) gait, like apes. Both have assumed that this bipedalism was almost exclusively on land, in trees or a combination of the two. Recent findings favoured the FU hypothesis by showing that the BK gait is 50–60% more energetically costly than a FU human gait on land. We confirm these findings but show that in water this cost differential is markedly reduced, especially in deeper water, at slower speeds and with greater knee flexion. These data suggest that the controversy about australopithecine locomotion may be eased if it is assumed that wading was a component of their locomotor repertoire and supports the idea that shallow water might have been an environment favourable to the evolution of early forms of “non-optimal” hominid bipedalism.     

An investigation into the usefulness of a cladistic approach to the study of the origin of anatomically modern humans

Cladistic analyses for the study of hominid evolution become very common during the last two decades, but little attention has been given to the appropriateness of the approach to studies being undertaken. This paper discusses how cladistic analyses have been used in studies of late Middle and Upper Pleicostocene hominids without due consideration of the problems inherent within the approach. It is concluded that in studies of the origin of anatomicaly modern humans a strict cladistic approach is inappropriate because it takes too narrow a view (presence/absence) of morphology, and in doing so does not allow for morphological variation. A phenetic approach which is interested in overall morphological similarity based on many characters and attempts to sample the total morphological variability evident within a sample would seem a more appropriate approach in such studies.     

Tool behavior and the origins of laterality

The cerebral cortex of modern humans is exceptional in that it is characterized by certain functional asymmetries for which no clear homologue is known in the non-human primates. These asymmetries consist at least in part in the presence in the left cerebral hemisphere of certain mechanisms which mediate language as well as skilled manual activity. Right-handedness is the most obvious overt manifestation of this cerebral asymmetry. It is argued in this paper that the lateralized representation of these mechanisms is an evolutionary consequence of the requirement for asymmetric employment of the forelimbs in the making and using of tools during hominid evolution. The adaptiveness of such an asymmetric arrangement is shown to follow from a few assumptions pertaining to brain organization and evolution. The colateralization of language mechanisms in modern humans to the left hemisphere is held to be a consequence of the coupling of these linguistic mechanisms to the motoric mechanisms already lateralized to the left hemisphere at an earlier point in hominid evolution. Finally, it is argued that this explanation of the evolution of laterality is more parsimonious in relation to known facts about human evolution than competing hypotheses.     

Evolution in metacommunities: on the relative importance of species sorting and monopolization in structuring communities

Abstract: Adaptive evolution within species and community assembly involving multiple species are both affected by dispersal and spatiotemporal environmental variation and may thus interact with each other. We examined this interaction in a simple three-patch metacommunity and found that these two processes produce very different associations between species composition and local environment. In most conditions, we find a pattern we call "species sorting," wherein local adaptation by resident species cannot prevent invasions by other preadapted species as environmental conditions change (strong association between local environmental conditions and local community composition). When dispersal rates are very low relative to the other two rates, local adaptation by resident species predominates, leading to strong priority effects that prevent successful colonization by other species that would have been well adapted, a pattern we call "local monopolization." When dispersal and evolutionary rates are both very high, we find that an evolving species outcompetes other species in all patches, a pattern we call "global monopolization." When environmental oscillations are very frequent, local monopolization predominates. Our findings indicate that there can be strong modification of community assembly by local adaptive processes and that these depend strongly on the relative rates of evolution, dispersal, and environmental change.     

Rates of evolution: Is there a conflict between neo-darwinian evolutionary theory and the fossil record?

Neo-darwinian and population genetics theory assumes that the necessary and sufficient set of conditions for all genetic, therefore evolutionary, change has been identified. Punctuationalists have assumed the opposite and cite the fossil record as evidence for change too rapid to be explained in neo-darwinian theory. Data is given here to provide estimates of the rate of evolution in hominid fossils, in living populations, and of that rate which would qualify as punctuational in the hominid fossil record. Evolution in living populations is orders of magnitude greater than that found in the fossil record and far greater than necessary to create apparently instantaneous saltations in the fossil record. It is suggested that such saltations may not represent more rapid rates of evolution but, rather, the persistence of evolutionary change in a given direction for a longer than normal period.     

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.     

French Contribution to the study of Human Origins: The case ofAustralopithecus afarensis

For a long time, French scientists have been involved in the study of human evolution and especially of human origins. Their key works in Eastern Africa have led to the discovery of major fossil hominid sites, especially in the Afar region in Ethiopia, where numerous remains ofAustralopithecus afarensis have been unearthed. The major contribution of the French scholars to the interpretation of the Hadar sample was to demonstrate the impact of the postcranial features on taxonomy and phylogeny. Two groups were identified in the sample and by comparison with modern populations of wild primates, these groups are assigned to different taxa. The other major impact was to show that early hominid bipedalism was an exact replica of modern human bipedality.     

Hominid paleoecology: The competitive exclusion principle and determinants of niche relationships

Some paleonanthropologists invoke the competitive exclusion principle programmatically in support of a single-species (or lineage) hypothesis of hominid evolution. Others apparently accept the association between competitive exclusion and a single species view, and develop multi-species interpretations using only taxonomic concepts. This paper demonstrates that the competitive exclusion principle itself is too assumption-bound to be appropriate for analyzing questions of hominid coexistence. Especially, the principle does not establish the ecological validity of the single-species hypothesis. Contrary to its immediate appearances, the principle has developed historically and analytically to predict and explain the evolution and maintenance of diversity in communities. Used in this manner it constitutes an important basis for the study of hominid paleoecology. Niche concepts necessary to demonstrate this are introduced, and competition and other factors that may have influenced the realized niches of sympatric hominid species are discussed. Nearly all modifications of the competitive exclusion principle that make it more realistic also illuminate factors that generate stable coexistence among competing species. These factors and their organizing theory establish the relevance of a broad data base to the analysis of hominid paleoecology; they should help to guide research on the ecology of early homonid species and their interrelationships.     

A new morphometric analysis of the hominid pelvic bone

This study is based upon a new morphometric technique providing both size and shape variables. It has been applied to 189 pelvic bones of extant humans and African apes as well as to 13 hominid pelvic bones of various taxonomic status. The main aim of this work is to include such fossil bones in the same study in order to set a synthetic comparison of their shape in the light of the yardstick given by the African ape/human pelvic bone comparison. To do so, ratio diagrams are chosen because they are simple and very expressive tools with which to present such comparisons. Shape differences are very well illustrated and quantified by this technique. The ilium appears to be the most different of the three parts of the pelvic bone. Compared to these differences, discrepancies between fossil hominid and extant human bones are of a totally different scale. This shows the architectural unity related to the acquisition of bipedalism by hominids. It is nonetheless possible to detect two levels of difference. The first separates Australopithecus from Homo and could be seen as reflecting locomotor differences between both genera. The second splits both Homo erectus and Neanderthal from modern human pelvic bones. It appears from the hominid fossil record of pelvic bones that two periods of stasis exist and are separated by a period of very rapid evolution corresponding to the emergence of the genus Homo. We are of the opinion that the same could be true for the split between African ape and hominid lineages at the end of the Miocene.     

An ecomorphological model of the initial hominid dispersal from Africa

We use new data on the timing and extent of the early Pleistocene dispersal of Homo erectus to estimate diffusion coefficients of early Homo from Africa. These diffusion coefficients indicate more rapid and efficient dispersals than those calculated for fossil Macaca sp., Theropithecus darti, and Mesopithecus pentelicus. Increases in home range size associated with changes in ecology, hominid body size, and possibly foraging strategy may underlay these differences in dispersal efficiency. Ecological data for extant primates and human foragers indicate a close relationship between body size, home range size, and diet quality. These data predict that evolutionary changes in body size and foraging behavior would have produced a 10-fold increase in the home range size of H. erectus compared with that of the australopithecines. These two independent datasets provide a means of quantifying aspects of the dispersal of early Homo and suggest that rapid rates of dispersal appear to have been promoted by changes in foraging strategy and body size in H. erectus facilitated by changes in ecosystem structure during the Plio-Pleistocene.     

History of American paleoanthropological research on early Hominidae, 1925–1980

Understanding of the early stages of hominid evolution prior to 1925 was based primarily on comparative morphological evidence derived from extant primates. With the publication of Australopithecus by Dart in 1925 and subsequent research in South Africa, new possibilities for empirical assessment of early hominid evolutionary history were opened. It was Gregory's work, with Hellman, reported at the first meeting of the AAPA in 1930, that convinced many workers of the hominid status of Australopithecus. The debunking of Eoanthropus as a Pliocene hominid, far from having a totally negative effect, showed that cranial expansion had occurred after bipedalism in hominid evolution, demonstrated that chemical dating had come of age, and in a broader sense, had underlined that phylogenetic hypotheses are falsifiable by recourse to the evidence. The input of biological sciences into early hominid studies, as exemplified by Washburn's “new physical anthropology,” reduced taxonomic diversity and focused attention on paleoecology and behavior. The development of the multidisciplinary approach to field research, pioneered by L. Leakey and brought to fruition by Howell, was of fundamental importance in accurately dating and understanding the context of early hominids. Archaeology, primatology, comparative and functional morphology, and morphometrics have contributed substantially in recent years to a fuller understanding of early hominid evolution. American granting agencies have heavily supported early hominid research but patterns of funding have not kept pace with the change from research based largely on individualistic enterprise to multidisciplinary research projects. Future early hominid research, if funding is available, will likely be directed toward investigating temporal and geographic gaps now known in the fossil record and in more rigorous and multidisciplinary investigations of early hominid behavior.