Fracture mechanics, enamel thickness and the evolution of molar form in hominins.

As the tissue most directly responsible for breaking down food in the oral cavity, the form and function of enamel is obviously of evolutionary significance in humans, non-human primates and other vertebrates. Accordingly, a standard metric, relative enamel thickness (RET), has been used for many decades to provide insights into vertebrate and human palaeobiology. Relatively thick enamel has evolved many times in vertebrates including hominoids (the group to which living humans and fossil hominins belong), and this pattern is thought to provide information about taxonomy, phylogeny, functional anatomy and diet. In particular, relatively thick enamel is thought to make tooth crowns strong so that they resist fractures associated with eating mechanically resistant foods. Here, we use current models of tooth biomechanics to show that RET is at best only moderately informative of function and diet in living hominoids and fossil hominins, and at worst provides misleading information. We propose a new metric, absolute crown strength, to assess the resistance of teeth to fracture, and identify what may be a novel characteristic of tooth strength in fossil hominins.

Assessing the reliability of virtual reconstruction of mandibles.

OBJECTIVES: Mandibular morphological variation is often used to examine various aspects of human palaeobiology. However, fossil and archeological skeletal remains are often fragmented/distorted and so are frequently excluded from studies. This leads to decreased sample sizes and, potentially, to biased results. Thus, it is of interest to restore the original anatomy of incomplete/distorted specimens. Thin plate splines (TPS), commonly used in Geometric Morphometrics (GM), offer the prospect of reconstruction of missing parts and particularly of interest here, missing landmarks. MATERIALS AND METHODS: Here, the reliability of TPS based mandibular reconstruction is tested. To that end missing landmarks were simulated in originally complete hemimandibles. TPS was then used to restore the location of simulated missing data and the predicted landmarks were compared to the original ones. RESULTS: Results show that error varies according to the number and location of estimated landmarks. Notwithstanding, estimation error is usually considerably smaller than the morphological differences between individuals from the same species. DISCUSSION: TPS based reconstruction allows fragmentary mandibles to be used in studies of whole mandibular variation, provided the above mentioned caveats are considered.

Last Common Ancestor of Apes and Humans: Morphology and Environment.

For much of their history, fossil apes retained many monkey-like features in posture and body structure. They also occupied a range of habitats, of which tropical forest was only a part, and there is evidence of increasing terrestriality in the fossil record as it is known at present (2019). In the early Miocene (18-20 million years ago, Ma), fossil apes were pronograde arboreal slow climbers, associated mainly with forest environments and deciduous woodland and with some indications of terrestrial behaviour, particularly the larger species. Their hands had long and opposable thumbs, and the phalanges were curved. In the early middle Miocene (15-16 Ma), apes were still monkey-like in body plan and posture and were associated almost entirely with non-forest, deciduous woodland habitats, with increasing evidence of terrestrial adaptations. Hand proportions remained the same. Towards the end of the middle Miocene (12 Ma), some fossil ape species had broadened chests, long clavicles, medial torsion of the humerus and re-positioning of the scapula to the back. These adaptations may have been linked with more upright posture, as in the living apes, but unlike them, the hand phalanges were short, robust and less curved, and the thumb remained long. Associated environments were deciduous woodland rather than forest. This body plan was retained in part in some later Miocene apes (10 Ma), some of which also had more elongated limbs and hands (thumb length not known), and hind limbs modified for greater flexibility, analogous with the orang utan. Associated environments were subtropical deciduous woodlands and subtropical evergreen laurophyllous woodland in southern Europe. Other late Miocene European apes had adaptations for living on the ground, and some of these also shared characters of the skull with orang utans. They are associated with more open deciduous woodland habitats. This body plan and environment were retained in the early hominin, Ardipithecus ramidus, but with a more robust postcranial skeleton and incipient bipedalism. Based on shared character states in fossil apes, living apes and early hominins, 27 characters are identified as probable attributes of the last common ancestor (LCA) of apes and humans. The likely environment of the LCA was tropical deciduous woodland with some evidence of more open habitats, and this remained unchanged in the transition from apes to early hominins.

Neanderthals as familiar strangers and the human spark: How the 'golden years' of Neanderthal research reopen the question of human uniqueness.

During the past decades, our image of Homo neanderthalensis has changed dramatically. Initially, Neanderthals were seen as primitive brutes. Increasingly, however, Neanderthals are regarded as basically human. New discoveries and technologies have led to an avalanche of data, and as a result of that it becomes increasingly difficult to pinpoint what the difference between modern humans and Neanderthals really is. And yet, the persistent quest for a minimal difference which separates them from us is still noticeable in Neanderthal research. Neanderthal discourse is a vantage point from which the logic of 'us' versus 'other' is critically reconsidered. Studying contemporary academic literature and science autobiographies from an oblique perspective, focusing not on Neanderthals as objects, but on the dynamics of interaction between Neanderthal researchers and their finds, basic convictions at work in this type of research are retrieved. What is at issue is not the actual distinction between modern humans and Neanderthals (which is continuously being redefined), but rather the dualistic construction of human and nonhuman. Neanderthal understanding is affected by the desire to safeguard human uniqueness. The overall trend is to identify the human mark or spark, which defines us as favoured 'winners'. The paradoxes emerging in contemporary Neanderthal discourse are symptomatic of the fact that a dualistic style of thinking is no longer tenable.

The biomechanical significance of the frontal sinus in Kabwe 1 (Homo heidelbergensis).

Paranasal sinuses are highly variable among living and fossil hominins and their function(s) are poorly understood. It has been argued they serve no particular function and are biological 'spandrels' arising as a structural consequence of changes in associated bones and/or soft tissue structures. In contrast, others have suggested that sinuses have one or more functions, in olfaction, respiration, thermoregulation, nitric oxide production, voice resonance, reduction of skull weight, and craniofacial biomechanics. Here we assess the extent to which the very large frontal sinus of Kabwe 1 impacts on the mechanical performance of the craniofacial skeleton during biting. It may be that the browridge is large and the sinus has large trabecular struts traversing it to compensate for the effect of a large sinus on the ability of the face to resist forces arising from biting. Alternatively, the large sinus may have no impact and be sited where strains that arise from biting would be very low. If the former is true, then infilling of the sinus would be expected to increase the ability of the skeleton to resist biting loads, while removing the struts might have the opposite effect. To these ends, finite element models with hollowed and infilled variants of the original sinus were created and loaded to simulate different bites. The deformations arising due to loading were then compared among different models and bites by contrasting the strain vectors arising during identical biting tasks. It was found that the frontal bone experiences very low strains and that infilling or hollowing of the sinus has little effect on strains over the cranial surface, with small effects over the frontal bone. The material used to infill the sinus experienced very low strains. This is consistent with the idea that frontal sinus morphogenesis is influenced by the strain field experienced by this region such that it comes to lie entirely within a region of the cranium that would otherwise experience low strains. This has implications for understanding why sinuses vary among hominin fossils.

Laetoli footprints reveal bipedal gait biomechanics different from those of modern humans and chimpanzees.

Bipedalism is a key adaptation that shaped human evolution, yet the timing and nature of its evolution remain unclear. Here we use new experimentally based approaches to investigate the locomotor mechanics preserved by the famous Pliocene hominin footprints from Laetoli, Tanzania. We conducted footprint formation experiments with habitually barefoot humans and with chimpanzees to quantitatively compare their footprints to those preserved at Laetoli. Our results show that the Laetoli footprints are morphologically distinct from those of both chimpanzees and habitually barefoot modern humans. By analysing biomechanical data that were collected during the human experiments we, for the first time, directly link differences between the Laetoli and modern human footprints to specific biomechanical variables. We find that the Laetoli hominin probably used a more flexed limb posture at foot strike than modern humans when walking bipedally. The Laetoli footprints provide a clear snapshot of an early hominin bipedal gait that probably involved a limb posture that was slightly but significantly different from our own, and these data support the hypothesis that important evolutionary changes to hominin bipedalism occurred within the past 3.66 Myr.

Reticulate evolution and the human past: an anthropological perspective.

CONTEXT: The evidence is mounting that reticulate (web-like) evolution has shaped the biological histories of many macroscopic plants and animals, including non-human primates closely related to Homo sapiens, but the implications of this non-hierarchical evolution for anthropological enquiry are not yet fully understood. When they are understood, the result may be a paradigm shift in evolutionary anthropology. OBJECTIVE/METHODS: This paper reviews the evidence for reticulated evolution in the non-human primates and human lineage. Then it makes the case for extrapolating this sort of patterning to Homo sapiens and other hominins and explores the implications this would have for research design, method and understandings of evolution in anthropology. RESULTS/CONCLUSION: Reticulation was significant in human evolutionary history and continues to influence societies today. Anthropologists and human scientists-whether working on ancient or modern populations-thus need to consider the implications of non-hierarchic evolution, particularly where molecular clocks, mathematical models and simplifying assumptions about evolutionary processes are used. This is not just a problem for palaeoanthropology. The simple fact of different mating systems among modern human groups, for example, may demand that more attention is paid to the potential for complexity in human genetic and cultural histories.

Topography of the mandibular canal in male human skulls originating from different time periods.

BACKGROUND: Dynamic advances in dentistry, especially in implantology has inspired researchers to carry out many studies investigating the topography of the mandibular canal and its ethnic differences. The aim of the study was a comparative analysis of variations in the position and topography of the mandibular canal based on radiographic images of human mandibles originating from modern and medieval skulls. MATERIALS AND METHODS: Morphometric examination of 126 radiographs of skulls (92 modern and 34 medieval skulls) was included. The age and sex of individuals were determined based on the morphology of the skull, the obliteration of cranial sutures, and the degree of tooth wear. To define the topography of the mandibular canal on X-ray images, we took 8 anthropometric measurements. RESULTS: We observed significant differences in several parameters. The distance between the base of the mandible and the bottom of the mandibular canal, the distance between the top of the mandibular canal and the crest of the alveolar arch, and the height of the mandibular body. Significant asymmetry was found for two parameters of mandibles from modern skulls: the distance between the top of the mandibular canal and the crest of the alveolar arch at the level of the second molar (p < 0.05), and the distance between the mandibular foramen and the margin of the anterior mandibular ramus (p < 0.007). There were no significant differences between measurements taken on the right and left sides of the medieval skulls. CONCLUSIONS: Our study revealed differences in the position of the mandibular canal between modern and medieval skulls, confirming the presence of geographical and chronological differences between populations. Knowledge of variability in the position of the mandibular canal between different local populations is fundamental for the correct interpretation of findings from diagnostic radiological studies used in dental practice and in forensic odontology or analysis of archaeological bone materials.

Topography of the infraorbital foramen in human skulls originating from different time periods.

BACKGROUND: The infraorbital foramen (IOF) is present on the maxilla under the infraorbital margin. Its identification is essential in various surgical procedures. The main aim of this study was the morphometric assessment of the position of the right and left infraorbital foramina in relation to specific structural elements of the facial skeleton, their width and direction, and also the determination of the location of these foramina above maxillary teeth in examined male skulls (belonging to European populations) dated to the beginning of the 20th century and the medieval and post-medieval period. This aim concerned also the assessment of the symmetry of the examined foramina (their location and size). An additional goal was to determine differences between the cranial samples concerning the analysed traits. MATERIALS AND METHODS: The six metric and two non-metric traits concerning the IOF were collected from the male cranial samples including modern skulls (n = 87), the medieval and post-medieval skulls (from 13th centuries and 15-17th centuries, respectively; n = 47) obtained from archaeological excavations in Wroclaw, and the sample of the medieval skulls (11-13th centuries, n = 100) from Sypniewo. The sex and age of the specimens were determined using the standard methodology. The appropriate statistical analysis was performed. RESULTS: Significant differences were established for three traits (taken from the left and right side) in the case of modern skulls (diameter of IOF, its distance to the midline, and zygomaticomaxillary suture) and one in the case of medieval skulls from Sypniewo (distance to the midline). In all of the cranial samples IOF most frequently occurred above the first upper molar. The greater diameter of IOF and its shorter distance to the alveolar crest and nasal notch were observed in non-modern skulls compared to modern skulls. CONCLUSIONS: The results of this study provide new additional data on the topography of IOF and its asymmetry, confirm the presence of both geographical and chronological differences between populations, and can be used in dental practice, and forensic odontology in the analysis of archaeological bone materials.

Integration of biplanar X-ray, three-dimensional animation and particle simulation reveals details of human 'track ontogeny'.

The emergence of bipedalism had profound effects on human evolutionary history, but the evolution of locomotor patterns within the hominin clade remains poorly understood. Fossil tracks record in vivo behaviours of extinct hominins, and they offer great potential to reveal locomotor patterns at various times and places across the human fossil record. However, there is no consensus on how to interpret anatomical or biomechanical patterns from tracks due to limited knowledge of the complex foot-substrate interactions through which they are produced. Here, we implement engineering-based methods to understand human track formation with the ultimate goal of unlocking invaluable information on hominin locomotion from fossil tracks. We first developed biplanar X-ray and three-dimensional animation techniques that permit visualization of subsurface foot motion as tracks are produced, and that allow for direct comparisons of foot kinematics to final track morphology. We then applied the discrete element method to accurately simulate the process of human track formation, allowing for direct study of human track ontogeny. This window lets us observe how specific anatomical and/or kinematic variables shape human track morphology, and it offers a new avenue for robust hypothesis testing in order to infer patterns of foot anatomy and motion from fossil hominin tracks.

Care of Infants in the Past: Bridging evolutionary anthropological and bioarchaeological approaches.

The importance of care of infants and children in palaeoanthropological and human behavioural ecological research on the evolution of our species is evident in the diversity of research on human development, alloparental care, and learning and social interaction. There has been a recent surge of interest in modelling the social implications of care provision for people with serious disabilities in bioarchaeology. However, there is a lack of acknowledgement of infant and child care in bioarchaeology, despite the significant labour and resources that are required, and the implications this has for health outcomes within societies. Drawing on the recent proliferation of studies on infancy and childhood in evolutionary anthropology and bioarchaeology, this paper presents ways the subdisciplines may draw on research developments from each field to advance a more holistic understanding of the evolutionary, social and health significance of infant and children care in the past.

Long-term patterns of body mass and stature evolution within the hominin lineage.

Body size is a central determinant of a species' biology and adaptive strategy, but the number of reliable estimates of hominin body mass and stature have been insufficient to determine long-term patterns and subtle interactions in these size components within our lineage. Here, we analyse 254 body mass and 204 stature estimates from a total of 311 hominin specimens dating from 4.4 Ma to the Holocene using multi-level chronological and taxonomic analytical categories. The results demonstrate complex temporal patterns of body size variation with phases of relative stasis intermitted by periods of rapid increases. The observed trajectories could result from punctuated increases at speciation events, but also differential proliferation of large-bodied taxa or the extinction of small-bodied populations. Combined taxonomic and temporal analyses show that in relation to australopithecines, early Homo is characterized by significantly larger average body mass and stature but retains considerable diversity, including small body sizes. Within later Homo, stature and body mass evolution follow different trajectories: average modern stature is maintained from ca 1.6 Ma, while consistently higher body masses are not established until the Middle Pleistocene at ca 0.5-0.4 Ma, likely caused by directional selection related to colonizing higher latitudes. Selection against small-bodied individuals (less than 40 kg; less than 140 cm) after 1.4 Ma is associated with a decrease in relative size variability in later Homo species compared with earlier Homo and australopithecines. The isolated small-bodied individuals of Homo naledi (ca 0.3 Ma) and Homo floresiensis (ca 100-60 ka) constitute important exceptions to these general patterns, adding further layers of complexity to the evolution of body size within the genus Homo. At the end of the Late Pleistocene and Holocene, body size in Homo sapiens declines on average, but also extends to lower limits not seen in comparable frequency since early Homo.

Virtual anthropology - a brief review of the literature and history of computed tomography.

Computed tomography (CT) has influenced numerous fields since its inception in the 1970s. The field of palaeoanthropology significantly benefited from this efficient and non-invasive medium in terms of the conservation, reconstruction and analysis of fossil human remains. Over the past decade, there has been a steady increase in the number of forensic anthropological studies incorporating virtual osteological analyses. Because of the increasing importance of these modern cross-sectional imaging techniques and the requirement for standardized parameters in forensic science, we deemed it important to outline the history and development of CT applications in these related academic areas. The present paper outlines the history of "virtual anthropology" and osteological multi-detector CT in the context of palaeoanthropology and forensic anthropology.

The discovery of fire by humans: a long and convoluted process.

Numbers of animal species react to the natural phenomenon of fire, but only humans have learnt to control it and to make it at will. Natural fires caused overwhelmingly by lightning are highly evident on many landscapes. Birds such as hawks, and some other predators, are alert to opportunities to catch animals including invertebrates disturbed by such fires and similar benefits are likely to underlie the first human involvements with fires. Early hominins would undoubtedly have been aware of such fires, as are savanna chimpanzees in the present. Rather than as an event, the discovery of fire use may be seen as a set of processes happening over the long term. Eventually, fire became embedded in human behaviour, so that it is involved in almost all advanced technologies. Fire has also influenced human biology, assisting in providing the high-quality diet which has fuelled the increase in brain size through the Pleistocene. Direct evidence of early fire in archaeology remains rare, but from 1.5 Ma onward surprising numbers of sites preserve some evidence of burnt material. By the Middle Pleistocene, recognizable hearths demonstrate a social and economic focus on many sites. The evidence of archaeological sites has to be evaluated against postulates of biological models such as the 'cooking hypothesis' or the 'social brain', and questions of social cooperation and the origins of language. Although much remains to be worked out, it is plain that fire control has had a major impact in the course of human evolution.This article is part of the themed issue 'The interaction of fire and mankind'.

The origins of the Acheulean: past and present perspectives on a major transition in human evolution.

The emergence of the Acheulean from the earlier Oldowan constitutes a major transition in human evolution, the theme of this special issue. This paper discusses the evidence for the origins of the Acheulean, a cornerstone in the history of human technology, from two perspectives; firstly, a review of the history of investigations on Acheulean research is presented. This approach introduces the evolution of theories throughout the development of the discipline, and reviews the way in which cumulative knowledge led to the prevalent explanatory framework for the emergence of the Acheulean. The second part presents the current state of the art in Acheulean origins research, and reviews the hard evidence for the appearance of this technology in Africa around 1.7 Ma, and its significance for the evolutionary history of Homo erectusThis article is part of the themed issue 'Major transitions in human evolution'.