Climatic influences on the evolution of early Homo?

The nature of the human fossil record is less than ideal for the generation of precise correlations between environmental variables and patterns of evolution in specific lineages. Nonetheless, a critical look at what can and cannot be said from individual fossil morphology and the correlation of specific environmental proxies with specific hominin fossils may lead to a greater understanding of the degree of certainty with which we should embrace environmental hypotheses for the evolution of Homo. Climate shifts have been implicated in both the origin of the genus and its dispersal from Africa. Here, I consider three areas in which a climatic influence has been posited to explain evolutionary shifts in the genus Homo: the origin and dispersal of the genus from Africa; geography, climate and body size in early Homo, and the influence of climate-induced sea level rise on morphological isolation in H. erectus. Each of the data sets is far from ideal, and interpretations of each of the data sets are fraught with issues of equifinality. Of the three hypotheses discussed, the clearest link is seen between latitudinal variation (and presumably temperature) and body size in H. erectus. Similarly, climate-induced sea level change seems a reasonable isolating mechanism to explain the pattern of cranial variation in later Asian H. erectus, but the distribution could also reflect incompletely sampled clinal variation. Alternatively, only equivocal support is found for the influence of climate on the differentiation of H. erectus from H. habilis (as proxied by body/brain size scaling), and therefore the dispersal of the genus Homo cannot be as clearly linked to changes in body size and shape as it has been in the past. These preliminary data suggest that an emphasis on understanding local adaptation before looking at global (and specific) level change is critical to elucidating the importance of climatic factors on the evolution of the genus Homo.     

The place of Neandertals in the evolution of hominid patterns of growth and development

This study uses the two developmental fields of dental maturation and femoral growth to determine if the pattern of growth and development in Neandertals (archaic Homo sapiens) was intermediate between that of Homo erectus and recent modern humans. Specimens used in the analysis included Neandertals and Upper Palaeolithic early modern Homo sapiens from Europe and individuals from two recent modern human populations. Ontogenetic data for the H. erectus adolescent KNM-WT 15000 and for Gorilla gorilla were included for comparison. Previous reports have indicated that H. erectus demonstrates a pattern of ontogeny characterized by earlier and more rapid linear growth than in modern humans. Results reported here demonstrate that Upper Paleolithic early modern Homo sapiens display a growth trajectory indistinguishable from that of recent modern humans. The pattern of Neandertal ontogeny is not intermediate between the pattern displayed in H. erectus and the derived pattern seen in the modern reference samples and the early modern H. sapiens sample. The Neandertal growth trajectory is consistent with either slow linear growth or advanced dental development.     

Brain size and encephalization in early to Mid-Pleistocene Homo

Important changes in the brain have occurred during the course of human evolution. Both absolute and relative size increases can be documented for species of Homo, culminating in the appearance of modern humans. One species that is particularly well-represented by fossil crania is Homo erectus. The mean capacity for 30 individuals is 973 cm(3). Within this group there is substantial variation, but brain size increases slightly in specimens from later time periods. Other Middle Pleistocene crania differ from those of Homo erectus. Characters of the facial skeleton, vault, and cranial base suggest that fossils from sites such as Arago Cave in France, the Sima de los Huesos in Spain, Bodo in Ethiopia, Broken Hill in Zambia, and perhaps Dali in China belong to the taxon Homo heidelbergensis. Ten of these mid-Quaternary hominins have brains averaging 1,206 cm(3) in volume, and many fall beyond the limits of size predicted for Homo erectus of equivalent age. When orbit height is used to construct an index of relative brain size, it is apparent that the (significant) increase in volume documented for the Middle Pleistocene individuals is not simply a consequence of larger body mass. Encephalization quotient values confirm this finding. These changes in absolute and relative brain size can be taken as further corroborative evidence for a speciation event, in which Homo erectus produced a daughter lineage. It is probable that Homo heidelbergensis originated in Africa or western Eurasia and then ranged widely across the Old World. Archaeological traces indicate that these populations differed in their technology and behavior from earlier hominins.     

Cranial capacity evolution in Homo erectus and early Homo sapiens.

This paper investigates patterns of cranial capacity evolution in Homo erectus, early Homo sapiens, and in regional subsamples of H. erectus. Specifically, models explaining evolution of cranial capacity in these taxa are evaluated with statistical techniques developed for the analysis of time series data. Regression estimates of rates of evolution in cranial capacity are also obtained. A non-parametric test for trend suggests that cranial capacity in both H. erectus and early H. sapiens may increase significantly through time. Cranial capacity in an Asian subsample of H. erectus (comprised of Chinese and Indonesian specimens) increases significantly through time. Other subsamples of H. erectus (African, Chinese, and Indonesian) do not appear to increase significantly through time. Regression results generally corroborate results of the test for trend. Spatial and temporal variation may characterize evolution of cranial capacity in H. erectus. Different patterns of cranial capacity evolution may distinguish H. erectus from early H. sapiens.     

中国古人类牙齿尺寸演化特点及东亚直立人的系统地位

对中国境内不同时代人类牙齿测量数据的发析显示,中国古人类牙齿尺寸的总体演化趋势与世界其它地区人类一致,呈缩小变化,其中一个表现特点是中国直立人与早期智人在牙齿尺寸上不差别不大,但作者根据对中国古人类化石形态特征,生存年代等方面的综合分析认为中国直立人在化石形态,生存年代等方面均与智人有明显的不同,取消直立人,将其并入智人意见的证据还是不够充分的,目前仍宜将直立人与智人作为人属内两个不同的种来看待,     

Early evidence of the genus Homo in East Asia

The timing and route of the earliest dispersal from Africa to Eastern Asia are contentious topics in the study of early human evolution because Asian hominin fossil sites with precise age constraints are very limited. Here we report new high-resolution magnetostratigraphic results that place stringent age controls on excavated hominin incisors and stone tools from the Yuanmou Basin, southwest China. The hominin-bearing layer resides in a reverse polarity magnetozone just above the upper boundary of the Olduvai subchron, yielding an estimated age of 1.7Ma. The finding represents the age of the earliest documented presence of Homo, with affinities to Homo erectus, in mainland East Asia. This age estimate is roughly the same as for H. erectus in island Southeast Asia and immediately prior to the oldest archaeological evidence in northeast Asia. Mammalian fauna and pollen obtained directly from the hominin site indicate that the Yuanmou hominins lived in a varied habitat of open vegetation with patches of bushland and forest on an alluvial fan close to a lake or swamp. The age and location are consistent with a rapid southern migration route of initial hominin populations into Eastern Asia.     

Cranial morphology of Javanese Homo erectus: new evidence for continuous evolution, specialization, and terminal extinction

Our current knowledge of the evolution of Homo during the early to middle Pleistocene is far from complete. This is not only because of the small number of fossil samples available, but also due to the scarcity of standardized datasets which are reliable in terms of landmark identification, interobserver error, and other distorting factors. This study aims to accurately describe the cranial morphological changes of H. erectus in Java using a standardized set of measurements taken by the authors from 18 adult crania from Sangiran, Trinil, Sambungmacan, and Ngandong. The identification of some obscure landmarks was aided by the use of micro-CT imaging. While recent studies tend to emphasize evolutionary conservatism in Javanese H. erectus, our results reinforce the theory that chronologically later groups experienced distinct morphological changes in a number of cranial traits. Some of these changes, particularly those related to brain size expansion, are similar to those observed for the genus Homo as a whole, whereas others are apparently unique specializations restricted to Javanese H. erectus. Such morphological specializations in Java include previously undescribed anteroposterior lengthening of the midcranial base and an anterior shift of the posterior temporal muscle, which might have influenced the morphology of the angular torus and supramastoid sulcus. Analyses of morphological variation indicate that the three crania from Sambungmacan variously fill the morphological gap between the chronologically earlier (Bapang-AG, Bapang Formation above the Grenzbank zone in Sangiran) and later (Ngandong) morphotypes of Java. At least one of the Bapang-AG crania, Sangiran 17, also exhibits a few characteristics which potentially indicate evolution toward the Ngandong condition. These strongly suggest the continuous, gradual morphological evolution of Javanese H. erectus from the Bapang-AG to Ngandong periods. The development of some unique features in later Javanese H. erectus supports the hypothesis that this Javanese lineage went extinct without making significant contributions to the ancestry of modern humans.     

Grandmothering and the evolution of homo erectus

Despite recent, compelling challenge, the evolution of Homo erectus is still commonly attributed to big game hunting and/or scavenging and family provisioning by men. Here we use a version of the "grandmother" hypothesis to develop an alternative scenario, that climate-driven adjustments in female foraging and food sharing practices, possibly involving tubers, favored significant changes in ancestral life history, morphology, and ecology leading to the appearance, spread and persistence of H. erectus. Available paleoclimatic, environmental, fossil and archaeological data are consistent with this proposition; avenues for further critical research are readily identified. This argument has important implications for widely-held ideas about the recent evolution of long human lifespans, the prevalence of male philopatry among ancestral hominids, and the catalytic role of big game hunting and scavenging in early human evolution.     

Dental microwear and diets of African early Homo

Conventional wisdom ties the origin and early evolution of the genus Homo to environmental changes that occurred near the end of the Pliocene. The basic idea is that changing habitats led to new diets emphasizing savanna resources, such as herd mammals or underground storage organs. Fossil teeth provide the most direct evidence available for evaluating this theory. In this paper, we present a comprehensive study of dental microwear in Plio-Pleistocene Homo from Africa. We examined all available cheek teeth from Ethiopia, Kenya, Tanzania, Malawi, and South Africa and found 18 that preserved antemortem microwear. Microwear features were measured and compared for these specimens and a baseline series of five extant primate species (Cebus apella, Gorilla gorilla, Lophocebus albigena, Pan troglodytes, and Papio ursinus) and two protohistoric human foraging groups (Aleut and Arikara) with documented differences in diet and subsistence strategies. Results confirmed that dental microwear reflects diet, such that hard-object specialists tend to have more large microwear pits, whereas tough food eaters usually have more striations and smaller microwear features. Early Homo specimens clustered with baseline groups that do not prefer fracture resistant foods. Still, Homo erectus and individuals from Swartkrans Member 1 had more small pits than Homo habilis and specimens from Sterkfontein Member 5C. These results suggest that none of the early Homo groups specialized on very hard or tough foods, but that H. erectus and Swartkrans Member 1 individuals ate, at least occasionally, more brittle or tough items than other fossil hominins studied.     

鄂西“南方古猿”和印尼早更新世若干人类化石

在鄂西发现的四枚臼齿化石曾被认为是南方古猿的。鄂西臼齿,从其齿冠尺寸和形状等来看,与非洲的有关材料对比,更接近人属成员的;与印尼早更新世有关的化石对比,与魁人等的很相似。直立人牙齿的演化趋势和变异性表明:鄂西臼齿以及印尼早更新世人类下颌骨化石更大的可能是代表一类时代较早的直立人。     

Middle Pleistocene human cranium from Tangshan (Nanjing), Southeast China: a new reconstruction and comparisons with Homo erectus from Eurasia and Africa

The morphology and affinities of early and middle Pleistocene Homo erectus in East Asia have been explored since the late nineteenth century. A fragmentary hominid cranium (Nanjing no.1) recovered in Tangshan near Nanjing, China bears directly on these issues. In the present study, the morphological features of Nanjing no.1 are described and compared with Homo erectus from both Eurasia and Africa. Our results indicate that this middle Pleistocene hominid fossil should be referred to as Homo erectus. The sharing of typical Homo erectus features with African and European counterparts demonstrates that Homo erectus is a widely distributed lineage that evolved during the million years after its Pliocene origins. The differences between Nanjing no.1 and Zhoukoudian suggest certain level of regional variation in East Asian Homo erectus.     

中国直立人与早期智人的牙齿形态鉴别

对直立人与早期智人的上颌牙齿特征的比较表明 :直立人牙齿的长宽尺寸 ,除中门齿唇舌径外 ,与早期智人的相应值很难区分 -早期智人的长宽尺寸多在直立人相应值的变异范围之内 ;直立人牙齿的观察性特征几乎都能在早期智人某些成员中见到。这意味着直立人与早期智人可能并无“种”而只有“亚种”这一分类级别上的差异 ,把直立人并入智人种这一建议是可取的。对若干化石地点的单个牙齿进行的重新鉴定表明 :桐梓、沂源、郧县梅铺、洛南和淅川的人类牙齿不一定是代表直立人的 ,有可能是代表早期智人的。     

The taxonomic implications of cranial shape variation in Homo erectus

The taxonomic status of Homo erectus sensu lato has been a source of debate since the early 1980s, when a series of publications suggested that the early African fossils may represent a separate species, H. ergaster. To gain further resolution regarding this debate, 3D geometric morphometric data were used to quantify overall shape variation in the cranial vault within H. erectus using a new metric, the sum of squared pairwise Procrustes distances (SSD). Bootstrapping methods were used to compare the H. erectus SSD to a broad range of human and nonhuman primate samples in order to ascertain whether variation in H. erectus most clearly resembles that seen in one or more species. The reference taxa included relevant phylogenetic, ecological, and temporal analogs including humans, apes, and both extant and extinct papionin monkeys. The mean cranial shapes of different temporogeographic subsets of H. erectus fossils were then tested for significance using exact randomization tests and compared to the distances between regional groups of modern humans and subspecies/species of the ape and papionin monkey taxa. To gauge the influence of sexual dimorphism on levels of variation, comparisons were also made between the mean cranial shapes of single-sex samples for the reference taxa. Results indicate that variation in H. erectus is most comparable to single species of papionin monkeys and the genus Pan, which included two species. However, H. erectus encompasses a limited range of variation given its extensive geographic and temporal range, leading to the conclusion that only one species should be recognized. In addition, there are significant differences between the African/Georgian and Asian H. erectus samples, but not between H. ergaster (Georgia+Africa, excluding OH 9 and Daka) and H. erectus sensu stricto. This finding is in line with expectations for intraspecific variation in a long-lived species with a wide, but probably discontinuous, geographic distribution.     

中国直立人形态特征的变异——颅骨测量数据的统计分析

近年,亚洲直立人化石特征的变异及其意义引起了学术界较多的关注。不断有学者提出周口店直立人无论在非测量性特征,还是在测量特征方面都具有不同于印度尼西亚及非洲直立人的表现特点,有人甚至对将周口店直立人化石特征作为直立人的典型特征的观点提出了置疑。为进一步探讨这些有争议的问题,本文采用多变量统计分析方法对33件非洲和欧亚地区直立人颅骨测量数据进行了分析。我们发现亚洲大陆的周口店和南京直立人与生活在东南亚群岛的印度尼西亚直立人具有不同的颅骨测量特征。中国标本以较窄的前额部和枕部,以及宽阔的颅骨中部为特点。而印度尼西亚标本的额部和枕部的宽度与颅骨中部的宽度相对较为接近。非洲直立人在这些颅骨测量特征的表现上与印度尼西亚直立人接近,呈现出相对宽阔的额部和枕部。中国的和县直立人在颅骨测量特征的表现上与印度尼西亚和非洲直立人接近,而与周口店和南京直立人明显不同。基于这些发现,本文就直立人头骨测量特征的稳定性、周口店直立人颅骨特征的表现特点、东亚直立人的地区变异与时代变化、头骨形态与测量特征的对应性等问题进行了讨论。     

Craniofacial morphology of Homo floresiensis: description, taxonomic affinities, and evolutionary implication

This paper describes in detail the external morphology of LB1/1, the nearly complete and only known cranium of Homo floresiensis. Comparisons were made with a large sample of early groups of the genus Homo to assess primitive, derived, and unique craniofacial traits of LB1 and discuss its evolution. Principal cranial shape differences between H. floresiensis and Homo sapiens are also explored metrically. The LB1 specimen exhibits a marked reductive trend in its facial skeleton, which is comparable to the H. sapiens condition and is probably associated with reduced masticatory stresses. However, LB1 is craniometrically different from H. sapiens showing an extremely small overall cranial size, and the combination of a primitive low and anteriorly narrow vault shape, a relatively prognathic face, a rounded oval foramen that is greatly separated anteriorly from the carotid canal/jugular foramen, and a unique, tall orbital shape. Whereas the neurocranium of LB1 is as small as that of some Homo habilis specimens, it exhibits laterally expanded parietals, a weak suprameatal crest, a moderately flexed occipital, a marked facial reduction, and many other derived features that characterize post-habilis Homo. Other craniofacial characteristics of LB1 include, for example, a relatively narrow frontal squama with flattened right and left sides, a marked frontal keel, posteriorly divergent temporal lines, a posteriorly flexed anteromedial corner of the mandibular fossa, a bulbous lateral end of the supraorbital torus, and a forward protruding maxillary body with a distinct infraorbital sulcus. LB1 is most similar to early Javanese Homo erectus from Sangiran and Trinil in these and other aspects. We conclude that the craniofacial morphology of LB1 is consistent with the hypothesis that H. floresiensis evolved from early Javanese H. erectus with dramatic island dwarfism. However, further field discoveries of early hominin skeletal remains from Flores and detailed analyses of the finds are needed to understand the evolutionary history of this endemic hominin species.     

中国远古人类的进化

本文从中国人类化石的年代顺序、共同形态特征、渐进变化、形态的异样性、镶嵌性、与其他地区的基因交流和古文化证据等方面论证了中国人类进化以连续性为主,还与世界其他地区之间有渐增的基因交流。本文还附带讨论了近年云南发现的古猿化石和湖北发现的人科头骨化石。     

Subnasal morphological variation in fossil hominids: a reassessment based on new observations and recent developmental findings

Quantitative and qualitative assessments of subnasal morphology in fossil hominids yield distinct patterns which have been used both to sort robust from nonrobust australopithecine taxa and to distinguish individual species. Recently, new developmental models have been applied to hominoid subnasal morphological variation. These studies require that certain features of the fossil hominid subnasal region, in particular the topography of the nasal cavity entrance and details of vomeral morphology, be reevaluated. This study does so for the robust and nonrobust australopithecines, early Homo (H. habilis/H. rudolfensis), and African H. erectus. Results reaffirm an overall similarity of the nonrobust Australopithecus subnasal morphological pattern with that of the chimpanzee. They further indicate that a vomeral insertion above the nasal surface of the premaxilla should be added to the list of traits characteristic of the robust australopithecine subnasal morphological pattern. Finally, reassessment of subnasal morphology in the early Homo and H. erectus samples from Africa suggest that these two taxa share a similar subnasal morphological pattern. This pattern consists of a smooth nasal cavity entrance, a horizontal nasal sill whose anterior edge is demarcated by a strong nasal crest, and a well-developed horizontal spine at the posterior edge of the nasal sill. Although none of the African fossil Homo specimens preserve a vomer, indirect evidence suggests that it would have inserted above the nasal sill.     

陕西公王岭蓝田直立人内耳迷路的复原及形态特点

1964年在陕西公王岭发现的蓝田人头骨的形态比周口店直立人和印度尼西亚爪哇直立人原始,其厚重的骨壁及较小的脑量,落入了早期人属成员的变异范围。最新测年结果将蓝田人的生存年代从原先普遍接受的距今115万年提早到大约163万年前,接近能人和南方古猿生存年代变异范围的下限,蓝田人是迄今为止我国发现的有确定年代数据的最早的古人类化石。本文采用高分辨率CT技术对蓝田人的颞骨岩部进行了扫描,对骨性内耳迷路进行了3D虚拟复原,通过与和县直立人、欧洲古老型智人、早期人属成员、南方古猿非洲种、粗壮傍人和现代人内耳迷路的21项测量项目的对比和分析,结果显示蓝田人内耳迷路的测量数据与南方古猿非洲种最接近,其次为现代人和欧洲古老型智人,而与早期人属成员和粗壮傍人相差较大。主成分分析结果显示,蓝田人内耳迷路与早期人属成员、欧洲古老型智人、南方古猿非洲种及现代人都有重叠区域,距离最近的是南方古猿非洲种Sts 5,其次为和县直立人和南方古猿非洲种Sts 19,而与粗壮傍人距离较远。本文研究提供了中更新世中国古人类内耳迷路的形态数据,为进一步探讨蓝田人体质特征演化上的意义提供了参考资料。     

Relative limb strength and locomotion in Homo habilis

The Homo habilis OH 62 partial skeleton has played an important, although controversial role in interpretations of early Homo locomotor behavior. Past interpretive problems stemmed from uncertain bone length estimates and comparisons using external bone breadth proportions, which do not clearly distinguish between modern humans and apes. Here, true cross-sectional bone strength measurements of the OH 62 femur and humerus are compared with those of modern humans and chimpanzees, as well as two early H. erectus specimens-KNM-WT 15000 and KNM-ER 1808. The comparative sections include two locations in the femur and two in the humerus in order to encompass the range of possible section positions in the OH 62 specimens. For each combination of section locations, femoral to humeral strength proportions of OH 62 fall below the 95% confidence interval of modern humans, and for most comparisons, within the 95% confidence interval of chimpanzees. In contrast, the two H. erectus specimens both fall within or even above the modern human distributions. This indicates that load distribution between the limbs, and by implication, locomotor behavior, was significantly different in H. habilis from that of H. erectus and modern humans. When considered with other postcranial evidence, the most likely interpretation is that H. habilis, although bipedal when terrestrial, still engaged in frequent arboreal behavior, while H. erectus was a completely committed terrestrial biped. This adds to the evidence that H. habilis (sensu stricto) and H. erectus represent ecologically distinct, parallel lineages during the early Pleistocene.     

脑膜中动脉的形态变异及其在人类进化上的意义

对现代人、直立人、早期智人、晚期智人脑膜中动脉前支和后支压迹的形态的观察和对比发现：从直立人到现代人脑膜中动脉管径从粗到逐渐变细,分枝从简单到逐渐复杂,前支趋向于增大,后支趋向于缩小,呈现出一定的规律性,有地域性差异。作者认为脑膜中动脉的这种分布形式间接地反映了直立人向智人演化及现代人类形成过程中,大脑各部分比例的变化和功能的日益复杂对供血需求的影响。在研究颅骨与大脑之间的关系上及脑演化上,脑膜中动脉有一定的意义。