The Neanderthal face is not cold adapted

Many morphological features of the Pleistocene fossil hominin Homo neanderthalensis, including the reputed large size of its paranasal sinuses, have been interpreted as adaptations to extreme cold, as some Neanderthals lived in Europe during glacial periods. This interpretation of sinus evolution rests on two assumptions: that increased craniofacial pneumatization is an adaptation to lower ambient temperatures, and that Neanderthals have relatively large sinuses. Analysis of humans, other primates, and rodents, however, suggests that the first assumption is suspect; at least the maxillary sinus undergoes a significant reduction in volume in extreme cold, in both wild and laboratory conditions. The second assumption, that Neanderthal sinuses are large, extensive, or even ‘hyperpneumatized,’ has held sway since the first specimen was described and has been interpreted as the causal explanation for some of the distinctive aspects of Neanderthal facial form, but has never been evaluated with respect to scaling. To test the latter assumption, previously published measurements from two-dimensional (2D) X-rays and new three-dimensional (3D) data from computed tomography (CT) of Neanderthals and temperate-climate European Homo sapiens are regressed against cranial size to determine the relative size of their sinuses. The 2D data reveal a degree of craniofacial pneumatization in Neanderthals that is both commensurate with the size of the cranium and comparable in scale with that seen in temperate climate H. sapiens. The 3D analysis of CT data from a smaller sample supports this conclusion. These results suggest that the distinctive Neanderthal face cannot be interpreted as a direct result of increased pneumatization, nor is it likely to be an adaptation to resist cold stress; an alternative explanation is thus required.     

Shape of the craniofacial complex in 45,X females: cephalometric study

The shape of the craniofacial complex was analysed cephalometrically in sixty-four adult 45,X females (Turner syndrome) using lateral skull radiographs, and the subjects were compared with first-degree female relatives and control females. The results showed that 45,X females have marked changes in relatively few craniofacial areas compared to the controls. Most of the changes are located in the cranial base, so that the face is retrognathic. The mandible is short, whereas the maxilla is of normal length. The results support the view that the morphology of the cranial base is markedly affected in 45,X females, whereas most other craniofacial changes could be considered secondary to the cranial base abnormality. It is suggested that retarded cartilage growth may be a factor leading to the present findings.     

Beneficial effect of hot spring bathing on stress levels in Japanese macaques

The ability of animals to survive dramatic climates depends on their physiology, morphology and behaviour, but is often influenced by the configuration of their habitat. Along with autonomic responses, thermoregulatory behaviours, including postural adjustments, social aggregation, and use of trees for shelter, help individuals maintain homeostasis across climate variations. Japanese macaques (Macaca fuscata) are the world’s most northerly species of nonhuman primates and have adapted to extremely cold environments. Given that thermoregulatory stress can increase glucocorticoid concentrations in primates, we hypothesized that by using an available hot spring, Japanese macaques could gain protection against weather-induced cold stress during winter. We studied 12 adult female Japanese macaques living in Jigokudani Monkey Park, Japan, during the spring birth season (April to June) and winter mating season (October to December). We collected faecal samples for determination of faecal glucocorticoid (fGC) metabolite concentrations by enzyme immunoassay, as well as behavioural data to determine time spent in the hot springs, dominance rank, aggression rates, and affiliative behaviours. We used nonparametric statistics to examine seasonal changes in hot spring bathing, and the relationship between rank and air temperature on hot spring bathing. We used general linear mixed-effect models to examine factors impacting hormone concentrations. We found that Japanese macaques use hot spring bathing for thermoregulation during the winter. In the studied troop, the single hot spring is a restricted resource favoured by dominant females. High social rank had both costs and benefits: dominant females sustained high fGC levels, which were associated with high aggression rates in winter, but benefited by priority of access to the hot spring, which was associated with low fGC concentrations and therefore might help reduce energy expenditure and subsequent body heat loss. This unique habit of hot spring bathing by Japanese macaques illustrates how behavioural flexibility can help counter cold climate stress, with likely implications for reproduction and survival.     

Implantation of cranial base metallic markers in nonhuman primates.

Cranial base metallic markers are useful in growth and developmental research on the nonhuman primate model. Metallic implants aid in superimposing serial cephalometric roentgenograms in the study of craniofacial changes. They also enable measurement of linear and angular changes in the cranial base. The design of a special implant gun is described in detail. A suggested technique for placement of tantalum markers in the cranial base of nonhuman primates is discussed.     

福建漳平奇和洞发现的新石器时代早期人类头骨

2011年在福建漳平奇和洞发现的距今1万年左右的新石器时代早期人类遗骸"奇和洞III号",是迄今在福建地区发现的最早、最完整的古人类头骨,为探讨华南更新世晚期向全新世过渡阶段人类的体质特征及现代人群的形成与分化提供了重要的研究材料。本文对这件头骨进行了研究,奇和洞III号为35岁左右的男性个体,牙齿龋病严重,推测当时人类的经济模式主要以农耕为主。通过与更新世晚期柳江、山顶洞101号及14组新石器时代人类头骨的比较,发现奇和洞III号头骨兼有更新世晚期人类及新石器南、北方居民的混合体质特征:奇和洞III号头骨长而脑量大,似更新世晚期人类;其高而狭窄的面部、宽阔而低矮的鼻部,呈现出不同于南、北方人群的特殊体质特征。主成分分析显示,奇和洞III号与对比的新石器时代各组在头骨的测量数据上没有表现为明显的南、北地区间差异,但在头骨的测量指数或形状上存在时代和地区间的不同。本文研究为新旧石器过渡阶段人类体质特征的变异提供了进一步证据。     

Craniofacial variation,body size and ecological factors in aboriginal populations from central Patagonia (2000–200 years B.P.)

Previous studies have shown that ecological factors had a significant role in shaping the patterns of craniofacial variation among South American populations. Here, we evaluate whether temperature and diet contributed to facial diversification in small geographic areas. Facial size and shape of 9 osteological samples from central Patagonia (Argentina) were described using 2D landmarks and semilandmarks. Data on mean annual temperature, diet composition (δ¹³C and δ¹⁵N values) and femoral head maximum breadth, used as a proxy of body mass, were obtained for each sample. We then tested the association of body mass and the ecological variables with facial morphology using spatial regression techniques and a model selection approach. Akaike Information Criterion produced disparate results for both components of facial morphology. The best model for facial size included temperature and body mass proxy, and accounted for more than 80% of variation in size. Lower temperatures were related to larger facial sizes. Body mass was negatively associated with facial size and showed no relationship with the temperature. This suggests a relatively independent variation of cranial traits and body mass at the spatial scale studied here. Facial shape was not associated with the temperature or diet composition, contrasting with the patterns observed at larger spatial scales. Our results point out that the effect of climatic variables on cranial traits might be a source of morphological differentiation not only at large scales but also in small geographic areas, and that size and shape display a differential preservation of environmental signals.     

EMG of serratus anterior and trapezius in the chimpanzee: scapular rotators revisited.

The importance of arm-raising has been a major consideration in the functional interpretation of differences in shoulder morphology among species of nonhuman primates. Among the characters that have been associated with enhancement of the arm-raising mechanism in hominoid primates are the relative enlargement of cranial trapezius and caudal serratus anterior, as the main scapular rotators, as well as changes in scapular morphology associated with their improved leverage for scapular rotation. Yet in an EMG study of cranial trapezius and caudal serratus anterior function in the great apes, Tuttle and Basmajian (Yrbk. Phys. Anthropol. 20:491-497, 1977) found these muscles to be essentially inactive during arm-raising. Although Tuttle and Basmajian suggest that the cranial orientation of the glenoid fossa in apes has reduced the demand for scapular rotation during arm-raising, subsequent EMG studies on other primate species suggest that these muscles do play a significant role in arm motion during active locomotion. This paper presents a reexamination of muscle recruitment patterns for trapezius and caudal serratus anterior in the chimpanzee. All but the lowest parts of caudal serratus anterior were found to be highly active during arm-raising motions, justifying earlier morphological interpretations of differences in caudal serratus anterior development. The lowest digitations of this muscle, while inactive during arm-raising, displayed significant activity during suspensory postures and locomotion, presumably to control the tendency of the scapula to shift cranially relative to the rib cage. Cranial trapezius did not appear to be involved in arm-raising; instead, its recruitment was closely tied to head position.     

Climate and the evolution of brachycephalization

Significant associations of cranial shape for 82 ethnic groups and seven climate variables are described. Variation among current populations is partially attributed to cold adaptation throughout the Pleistocene. Application of data files tabulated by the authors is described for a number of problems. Temporal distribution of 115 specimens indicates a geometric trend (CI = 76.7-1.96 log time X 10(3) ). Cranial indices are summarized within alternative taxonomic models and between climatic ecotypes. Evidence supports the hypothesis of cold adaptation among "Classic" Neandertals. Limitations of the thermodynamic model are discussed. It is probable that a decrease of the cranial index occurs from the Middle to Upper Paleolithic. During the Holocene, the index increases under all climatic conditions.     

Were neandertal and modern human cranial differences produced by natural selection or genetic drift?

Most evolutionary explanations for cranial differences between Neandertals and modern humans emphasize adaptation by natural selection. Features of the crania of Neandertals could be adaptations to the glacial climate of Pleistocene Europe or to the high mechanical strains produced by habitually using the front teeth as tools, while those of modern humans could be adaptations for articulate speech production. A few researchers have proposed non-adaptive explanations. These stress that isolation between Neandertal and modern human populations would have lead to cranial diversification by genetic drift (chance changes in the frequencies of alleles at genetic loci contributing to variation in cranial morphology). Here we use a variety of statistical tests founded on explicit predictions from quantitative- and population-genetic theory to show that genetic drift can explain cranial differences between Neandertals and modern humans. These tests are based on thirty-seven standard cranial measurements from a sample of 2524 modern humans from 30 populations and 20 Neandertal fossils. As a further test, we compare our results for modern human cranial measurements with those for a genetic dataset consisting of 377 microsatellites typed for a sample of 1056 modern humans from 52 populations. We conclude that rather than requiring special adaptive accounts, Neandertal and modern human crania may simply represent two outcomes from a vast space of random evolutionary possibilities.     

MAMMALIAN EXTINCTIONS IN THE LATE PLEISTOCENE OF NORTHERN EURASIA AND NORTH AMERICA

The 'mass extinctions' at the end of the Pleistocene were unique, both in the Pleistocene and earlier in the geological record, in that the species lost were nearly all large terrestrial mammals. Although a global phenomenon, late Pleistocene extinctions were most severe in North America, South America and Australia, and moderate in northern Eurasia (Europe plus Soviet Asia). In Africa, where nearly all of the late Pleistocene 'megafauna' survives to the present day, losses were slight. Ruling out epidemic disease or cosmic catastrophe, the contending hypotheses to explain late Pleistocene extinctions are: (a) failure to adapt to climatic/environmental change; and (b) extermination by human hunters ('prehistoric overkill'). This review focuses on extinctions in northern Eurasia (mainly Europe) in comparison with North America. In addition to reviewing the faunal evidence, the highly relevant environmental and archaeological backgrounds are summarized. The latest survival dates of extinct species are estimated from stratigraphic occurrences of fossil remains, radiocarbon dates, or association with archaeological industries. The Middle and Upper Pleistocene (ca. 700,000-10,000 BP) in northern Eurasia and North America was a time of constantly changing climate, ranging from phases of extensive glaciation in cold stages, to temperate periods (interglacials). In the Lateglacial (ca. 15,000-10,000 BP), during which most extinctions occurred, there was a major reorganization of vegetation, mainly involving the replacement of open vegetation by forests. These changes were more profound than earlier in the Last Cold Stage, but similar in nature to vegetational changes that took place at previous cold stage/interglacial transitions. The archaeological record shows that humans have been present in Europe since the early Middle Pleistocene. The arrival in Europe ca. 35,000 BP of 'anatomically modern humans', with their technologically more advanced upper palaeolithic industries, was a 'quantum leap' in human history. Extinctions occurred throughout the European Pleistocene, but until the late Pleistocene most losses were replaced by the evolution or immigration of new species, and most of those lost without replacement were small mammals. In marked contrast, extinctions without replacement in the late Pleistocene were almost entirely confined to the largest mammals (greater than 1000 kg) and some medium-large species (100-1000 kg).(ABSTRACT TRUNCATED AT 400 WORDS)     

Head shape variation among cryptic populations of ground skinks (Scincella lateralis)

Recent interest in testing the river–refugia effect has prompted an evolutionary investigation of the ground skink (Scincella lateralis) to diagnose instances of the riverine barrier hypothesis and thus, Pleistocene climate change refugia. This taxon, characterized by limited vagility, presents itself as a model organism in understanding patterns in Gulf Coast squamate evolution and biogeography. Here, we use geometric morphometric techniques to assess whether changes in head shape characterize diverging molecular lineages. We analysed head shape variation for diagnostic morphology by population and hypothesized that clades recovered as monophyletic by previous molecular studies will be more similar to each other than distantly related lineages. Dorsal and lateral head shape analyses indicated strong divergence for one clade (mitochondrial Clade I) in association with mtDNA divergence, exhibiting cranial narrowing and elongation relative to all other clades, and weak divergence of another clade (mitochondrial Clade N), which exhibited similar shape divergence from consensus. Both of these clades are Gulf Coast lineages and our results suggest convergence in head shape towards a regional phenotype.     

三峡地区最晚更新世的梅氏犀兼述中国南方更新世的犀牛化石

重庆市巫山县迷宫洞是一处重要的晚更新世晚期的古人类遗址, 其绝对年代为距今13150±190年。本文研究了这一遗址出土的犀牛化石, 材料包括头骨碎块、下颌、牙齿及数件头后骨骼。这些材料被鉴定为梅氏犀或称基什贝尔格犀(Stephanorhinus kirchbergensis), 是这个种已知时代最晚和分布最为靠南的记录。适合温带气候的S.kirchbergensis在中-晚更新世南下至长江流域, 可能是受到冰期寒冷气候的压迫所致。以迷宫洞和其他长江流域的S.kirchbergensis为基础, 通过对比, 认为我国南方、北方以及欧洲的S.kirchbergensis在颊齿形态上存在明显的区别, 从而将这个种分为三个类型。另外, 本文还对我国南方更新世的犀牛种类, 特别是中国犀(Rhinoceros sinensis), 进行了全面地厘定。     

Cephalometric analysis of the craniofacial region of the northern Foxe Basin Eskimo

Past investigations of the Eskimo have indicated that there are marked morphological differences in the craniofacial skeleton of this relatively isolated ethnic group compared to other ethnic and racial groups. This study, using cephalometric radiography, attempted to characterize the craniofacial phenotype of the Eskimo living in the northern Foxe Basin, Northwest Territories, Canada. Age changes were examined on a cross-sectional basis with comparisons being made with a Winnipeg Caucasian group. This investigation indicates that the Igloolik Eskimo has a phenotype, established early in life, and is distinct from the Winnipeg group. The overall size of the Eskimo craniofacial complex was significantly larger at three years of age and remained larger through the ages studied. Development of the craniofacial region, however, was fairly similar in rate and direction for both populations. The greatest differences between the Eskimo and Caucasian groups were found in the linear measurements assessing cranial width, facial width, mandibular length, facial height, protrusion of the incisors, chin point development, and nasal morphology. Differences between the two groups in the morphological relationships of the component structures include the angular relationships of the maxilla and nasal bones to the anterior cranial base, the gonial angle of the mandible, and the angle of facial convexity.     

Postnatal Cranial Development in Papionin Primates: An Alternative Model for Hominin Evolutionary Development

The evolution of hominin growth and life history has long been a subject of intensive research, but it is only recently that paleoanthropologists have considered the ontogenetic basis of human morphological evolution. To date, most human EvoDevo studies have focused on developmental patterns in extant African apes and humans. However, the Old World monkey tribe Papionini, a diverse clade whose members resemble hominins in their ecology and population structure, has been proposed as an alternative model for human craniofacial evolution. This paper reviews prior studies of papionin development and socioecology and presents new analyses of juvenile shape variation and ontogeny to address fundamental questions concerning primate cranial development, including: (1) When are cranial shape differences between species established? (2) How do epigenetic influences modulate early-arising pattern differences? (3) How much do postnatal developmental trajectories vary? (4) What is the impact of developmental variation on adult cranial shape? and, (5) What role do environmental factors play in establishing adult cranial form? Results of this inquiry suggest that species differences in cranial morphology arise during prenatal or earliest postnatal development. This is true even for late-arising features that develop under the influence of epigenetic factors such as mechanical loading. Papionins largely retain a shared, ancestral pattern of ontogenetic shape change, but large size and sexual dimorphism are associated with divergent developmental trajectories, suggesting differences in cranial integration. Developmental simulation studies indicate that postnatal ontogenetic variation has a limited influence on adult cranial morphology, leaving early morphogenesis as the primary determinant of cranial shape. The ability of social factors to influence craniofacial development in Mandrillus suggests a possible role for phentotypic plasticity in the diversification of primate cranial form. The implications of these findings for taxonomic attribution of juvenile fossils, the developmental basis of early hominin characters, and hominin cranial diversity are discussed.     

Cranial thickening in an Australian hominid as a possible palaeoepidemiological indicator

This paper describes the cranial thickening of a late Pleistocene hominid (Willandra Lakes Hominid 50) from Australia. The unusual development of the vault structures in this individual has few, if any, equals among other hominids or more recent populations from around the world. The vault morphology is, therefore, described in terms of a pathologically related condition associated with the modern haemolytic blood dyscrasias, typical of sickle cell anamia and thalassemia. A possible palaeoepidemiology for these genetic adaptations among early Australasian populations is proposed together with a discussion of similar changes observed in the vault of the Singa calvarium from the Sudan. It is tentatively suggested that the cranial thickening of the Australian hominid has its origins in some form of genetic blood disease and that if this diagnosis is correct, this individual provides a rare glimpse of human biological adaptation in the late Upper Pleistocene.     

Anterior cervical microsurgical approach to the cranial base in the rabbit: technical note.

Current trends in research on craniofacial syndromes have led to enhanced interest in the cranial base as a contributory factor in the development of normal and abnormal midfacial structure. Indeed, attention has focused upon one particular growth plate in the posterior cranial base, the spheno-occipital synchondrosis, since it has been shown that alterations in this structure are associated with profound changes in craniofacial growth. In this report we describe a surgical approach to the cranial base of the rabbit that is safe, simple and reliable. It is applicable to neonatal as well as adult rabbits.     

The Developmental Basis of Quantitative Craniofacial Variation in Humans and Mice

The human skull is a complex and highly integrated structure that has long held the fascination of anthropologists and evolutionary biologists. Recent studies of the genetics of craniofacial variation reveal a very complex and multifactorial picture. These findings contrast with older ideas that posit much simpler developmental bases for variation in cranial morphology such as the growth of the brain or the growth of the chondrocranium relative to the dermatocranium. Such processes have been shown to have major effects on cranial morphology in mice. It is not known, however, whether they are relevant to explaining normal phenotypic variation in humans. To answer this question, we obtained vectors of shape change from mutant mouse models in which the developmental basis for the craniofacial phenotype is known to varying degrees, and compared these to a homologous dataset constructed from human crania obtained from a single population with a known genealogy. Our results show that the shape vectors associated with perturbations to chondrocranial growth, brain growth, and body size in mice do largely correspond to axes of covariation in humans. This finding supports the view that the developmental basis for craniofacial variation funnels down to a relatively small number of key developmental processes that are similar across mice and humans. Understanding these processes and how they influence craniofacial shape provides fundamental insights into the developmental basis for evolutionary change in the human skull as well as the developmental-genetic basis for normal phenotypic variation in craniofacial form.     

Asymmetric vault modification in Hopi crania

Cradleboarding was practiced by numerous prehistoric and historic populations, including the Hopi. In this group, one result of cra-dleboarding was bilateral or asymmetric flattening of the posterior occipital. We test whether cradleboarding had significant effects on the morphology of the cranial vault, cranial base, and face. Additionally, we examine associations between direction of flattening and asymmetric craniofacial growth. A skeletal sample of Hopi from the Old Walpi site includes both nonmodified (N = 43) and modified individuals (N = 39). Three-dimensional coordinates of 53 landmarks were obtained using a diagraph. Thirty-six landmarks were used to define nine finite elements in the cranial vault, cranial base, and face. Finite element scaling was used to compare average nonmodified individuals, with averages of bilaterally, right, and left modified individuals. The significance of variation among “treatment” groups was evaluated using a bootstrap test. Pearson product-moment correlations test the association of asymmetry with direction of modification. Hopi cradleboarding has a significant effect on growth of the cranial vault, but does not affect morphology of the cranial base or face. Bilateral flattening of the cranial vault leads to decreased length and increased width of the cranial vault. Flattening of the right or left cranial vault results in ipsilaterally decreased length and width coupled with a corresponding increased length and width on the contralateral side of the cranial vault. There is a significant correlation of size asymmetry with direction of modification in the cranial vault, but not with size or shape change in the cranial base or face. © 1995 Wiley-Liss, Inc.     

Craniofacial variability and modularity in macaques and mice

Evolutionary developmental biology of primates will be driven largely by the developmental biology of the house mouse. Inferences from how known developmental perturbations produce phenotypic effects in model organisms, such as mice, to how the same perturbations would affect craniofacial form in primates must be informed by comparisons of phenotypic variation and variability in mice and the primate species of interest. We use morphometric methods to compare patterns of cranial variability in homologous datasets obtained for two strains of laboratory mice and rhesus macaques. C57BL/6J represents a common genetic background for transgenic models. A/WySnJ mice exhibit altered facial morphology which results from reduction in the growth of the maxillary process during formation of the face. This is relevant to evolutionary changes in facial prognathism in nonhuman primate and human evolution. Rhesus macaques represent a nonhuman primate about which a great deal of phenotypic and genetic information is available. We find significant similarities in covariation patterns between the C57BL/6J mice and macaques. Among-trait variation in genetic and phenotypic variances are fairly concordant among the three groups, but among-trait variation in developmental stability is not. Finally, analysis of modularity based on phenotypic and genetic correlations did not reveal a consistent pattern in the three groups. We discuss the implications of these results for the study of evolutionary developmental biology of primates and outline a research strategy for integrating mouse genomics and developmental biology into this emerging field.     

The ontogeny of cranial base angulation in humans and chimpanzees and its implications for reconstructing pharyngeal dimensions

This paper examines differences in the processes by which the cranial base flexes in humans and extends in chimpanzees. In addition, we test the extent to which one can use comparisons of cranial base angles in humans and non-human primates to predict vocal tract dimensions. Four internal cranial base angles and one external cranial base angle were measured in a longitudinal sample of Homo sapiens and a cross-sectional sample of Pan troglodytes. These data show that the processes of cranial base angulation differ substantially in these species. While the human cranial base flexes postnatally in a rapid growth trajectory that is complete by two years, the cranial base in P. troglodytes extends postnatally in a more prolonged skeletal growth trajectory. These comparisons also demonstrate that the rate of cranial base angulation is comparable for different measures, but that angles which incorporate different anterior cranial base measurements correlate poorly. We also examined ontogenetic relationships between internal and external cranial base angles and vocal tract growth in humans to test the hypothesis that cranial base angulation influences pharyngeal dimensions and can, therefore, be used to estimate vocal tract proportions in fossil hominids. Our results indicate that internal and external cranial base angles are independent of the horizontal and vertical dimensions of the vocal tract. Instead, a combination of mandibular and palatal landmarks can be used to predict dimensions of the vocal tract in H. sapiens. The developmental contrasts in cranial base angulation between humans and non-human primates may have important implications for testing hypotheses about the relationship between cranial base flexion and other craniofacial dimensions in hominid evolution.