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.     

Skeletal age, dental age, and the maturation of KNM-WT 15000

The skeleton of the Homo erectus boy from West Lake Turkana, Kenya (KNM-WT 15000), is remarkably complete, and this individual has thus provided a case study for several researchers examining Homo erectus growth. Using data from a longitudinal study of Montreal French-Canadian children, it is shown that while dental and skeletal ages match reasonably well at the level of a sample of children, individuals can display differences between skeletal and dental ages of 2 years or more. Furthermore, the relationship between these two markers may change over time in individual children. It is also possible to find children with patterns of dental maturation similar to KNM-WT 15000's pattern in the Montreal sample. Therefore, neither the discrepancy between skeletal age and dental age alone nor the pattern of dental maturation as assessed by dental stages precludes a human-like pattern of growth, including an adolescent growth spurt, for this individual. Some indicators (e.g., estimated body size for predicted age, and enamel formation) do suggest possible growth-patterning differences from modern humans, and therefore earlier maturation is a reasonable hypothesis, but caution is warranted, given the large degree of modern human variation in developmental markers and the inherent uncertainty in precise estimation of KNM-WT 15000's maturational parameters.     

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,     

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。     

中国直立人牙齿特征变异及其演化意义

以往研究发现,中国直立人化石呈现较大的形态变异。对于这种变异程度及造成变异的原因,在古人类学界有不同的认识。有学者提出在直立人阶段,中国古人类已经呈现明显的区域性差异,但也有人认为这些差异似乎没有规律性。近年,本文作者采用不同方法对若干地点的中国直立人牙齿特征及其变异进行了系列研究,取得了一些新的发现和认识。本文在回顾总结这些研究的基础上,结合对其它一些地点中国直立人牙齿特征的观测对比,对中国直立人牙齿特征表现特点及变异作了进一步的分析。本研究发现,中国直立人牙齿特征具有较大的变异范围。这些变异似可分为两种主要类型,元谋、建始、郧县梅铺、和县牙齿呈现出较多的原始特点,代表一种原始类型;周口店、沂源等地点的标本特征相对进步,表现出更多的典型直立人特征。其它一些地点的直立人化石呈现出混合或中间状态。值得注意的是部分呈现出原始牙齿特征类型的中国直立人的生存年代相对较晚,其牙齿特征的原始性与生存时代不具有对应关系。作者认为中国直立人牙齿特征类型反映了更新世早期和中期不同直立人群的演化状态。原始类型牙齿特征的形成不仅与演化时序性和地理分布有关,还在一定程度上反映了一些中国直立人群的演化隔离。     

Endocranial cast of Hexian Homo erectus from South China

In this paper, we present data on the morphological features and linear measurements for the Hexian Homo erectus and other comparative endocasts, in order to highlight variation during human brain evolution. The endocast of Hexian was reconstructed in 1982, and an endocranial volume of 1,025 ml was estimated. The geological age is about 412 ka, or roughly contemporaneous with the Zhoukoudian (ZKD) specimens. There are some differences between Hexian and the modern Chinese male endocasts in our sample, including low position of the greatest breadth, low maximum height, a well-marked and prominent frontal keel, the flat surface of the frontal lobes, prominent sagittal keel along the center frontal and parietal lobes, depressed Sylvian areas and parietal lobes superiorly, strong posterior projection of the occipital lobes, anterior position of the cerebellar lobes relative to the occipital lobes, and the relative simplicity of the meningeal vessels. Compared with the ZKD, Indonesian, and African Homo erectus specimens, Hexian has more morphological features in common with ZKD. Principal component analyses indicate that Hexian is closest to the ZKD Homo erectus compared with the modern Chinese and other Homo erectus, but its great breadth distinguishes it. Metric analyses show that the brain height, frontal breadth, cerebral height, frontal height, and parietal chord from Homo erectus to modern humans increased, while the length, breadth, frontal chord, and occipital breadth did not change substantially.     

中更新世非洲Bodo人类头骨化石与周口店直立人的比较--中国与非洲人类头骨特征对比之三

发现于埃塞俄比亚MiddleAwash地区Bodo地点距今60万年的人类头骨化石是迄今发现的最为古老和完整的非洲中更新世人类化石。由于Bodo头骨化石在形态特征上兼有直立人与智人的特点,多年来学术界对其分类地位一直存在争议。Rightmire认为Bodo头骨化石与BrokenHill及Petralona等在分类上属于古老型智人的中更新世人类更为接近,是非洲直立人向古老型智人过渡的代表。至少在距今60万年的中更新世早期直立人向古老型智人转变的成种事件在非洲就已经发生。以Bodo头骨为代表的一批更新世中期非洲和欧洲人类化石构成了可能是后期人类祖先的人属海德堡种。这些观点导致了近年学术界对古老型智人在非洲及欧亚出现时间以及更新世中期非洲和欧亚地区古人类相互之间演化关系的关注。基于这样的背景,本文对年代与Bodo化石接近的周口店直立人头骨特征与Bodo头骨的相似及差异表现情况进行了对比研究。结果发现Bodo头骨在一系列特征上与周口店直立人相似,同时在包括颅容量在内的其它一些特征上呈现出后期智人的特点,但总体形态上似乎与直立人更为相似。作者认为尽管这种进化上的镶嵌现象在中国古人类化石记录上也广泛存在,但由于中国人类化石标本在年代上的不确定性,目前还没有可靠的证据说明这种集直立人与智人化石特征为一体的镶嵌性在中国古人类化石出现的时间接近或早于非洲。考虑到中国与非洲直立人生存年代的巨大差异及人类演化的不同步或地区间差异,具有较多后期人类特征表现的人类首先出现在非洲是完全可能的。根据这些研究对比,作者就人类演化的镶嵌现象、更新世中期非洲与亚洲地区人类演化上的差异等问题进行了讨论。     

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。     

南京1号与东非Bodo人类头骨化石:对"中心和边缘"假说的检测

"中心和边缘"假说认为非洲是人类演化的中心地区,东亚等地区是边缘地区。在边缘地区,人群的地区性形态特征出现较早,可上溯到直立人生活时期;在中心地区,人群的地区性形态特征出现较晚。Bodo人类头骨化石和南京1号人类头骨化石分别出自中心地区和边缘地区,二者年代都是距今60万年左右,二者都保留有面颅。因此,Bodo人类头骨化石和南京1号人类头骨化石是检测"中心和边缘"假说的最合适的材料。本文是对南京1号和Bodo头骨的面颅测量性特征作比较研究。研究结果表明:1.二者面颅测量性特征上的差别远大于这两个相应地区现代人群之间的差别,提示了人类的地区性体质形态差别早在60万年前就很明显;2.东亚的南京1号人类头骨和东非的Bodo人类头骨尽管同样古老,但各自与当地区的现代人群的面颅上的差异情况并不一致。Bodo头骨与东非现代人群显得差异较大,南京1号头骨与东亚现代人群显得较相近。这种相近,提示了在东亚这个"边缘地区",现代人群的面颅测量性特征可追溯到以南京1号头骨为代表的远古人类那里,而在"中心地区",现代人群的面颅测量性特征还很难与以Bodo为代表的远古人群相联系。本项研究结果与"中心和边缘"假说的推测相符合。     

中国早期智人牙齿化石

中国目前较为可靠的早期智人牙齿材料应包括在巢县、许家窑、长阳、周口店新洞和桐梓发现的人类牙齿化石,代表一类与早期组早期智人相当或十分相近的古人类。丁村人类牙齿化石可能是代表晚期组早期智人也可能是代表解剖学上的现代智人。就牙齿材料而言,尚无充分的证据能表明在中国曾同时存在过两种类型的早期智人。     

Femoral/humeral strength in early African Homo erectus

Lower-to-upper limb-bone proportions give valuable clues to locomotor behavior in fossil taxa. However, to date only external linear dimensions have been included in such analyses of early hominins. In this study, cross-sectional measures of femoral and humeral diaphyseal strength are determined for the two most complete early Homo erectus (or ergaster) associated skeletons--the juvenile KNM-WT 15000 and the adult KNM-ER 1808. Modern comparative samples include an adult human skeletal sample representative of diverse body shapes, a human longitudinal growth series, and an adult chimpanzee sample. When compared to appropriately age-matched samples, both H. erectus specimens fall very close to modern human mean proportions and far from chimpanzee proportions (which do not overlap with those of humans). This implies very similar mechanical load-sharing between the lower and upper limbs, and by implication, similar locomotor behavior in early H. erectus and modern humans. Thus, by the earliest Pleistocene (1.7 Ma), completely modern patterns of bipedal behavior were fully established in at least one early hominin taxon.     

Homo floresiensis and the evolution of the hominin shoulder

The holotype of Homo floresiensis, diminutive hominins with tiny brains living until 12,000 years ago on the island of Flores, is a partial skeleton (LB1) that includes a partial clavicle (LB1/5) and a nearly complete right humerus (LB1/50). Although the humerus appears fairly modern in most regards, it is remarkable in displaying only 110 degrees of humeral torsion, well below modern human average values. Assuming a modern human shoulder configuration, such a low degree of humeral torsion would result in a lateral set to the elbow. Such an elbow joint would function more nearly in a frontal than in a sagittal plane, and this is certainly not what anyone would have predicted for a tool-making Pleistocene hominin. We argue that Homo floresiensis probably did not have a modern human shoulder configuration: the clavicle was relatively short, and we suggest that the scapula was more protracted, resulting in a glenoid fossa that faced anteriorly rather than laterally. A posteriorly directed humeral head was therefore appropriate for maintaining a normally functioning elbow joint. Similar morphology in the Homo erectus Nariokotome boy (KNM-WT 15000) suggests that this shoulder configuration may represent a transitional stage in pectoral girdle evolution in the human lineage.     

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。     

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.     

更新世中期中国古人类演化区域连续性与多样性的化石证据

以往,在东亚大陆发现的更新世中期人类化石被分别归入直立人和古老型智人。这种分类的主要依据是化石形态特征以及年代。魏敦瑞对周口店第一地点人类化石研究描述的一些头骨、下颌骨和牙齿特征通常被作为判定直立人的标准。根据这些化石的年代分布,一般将30万年前的中更新世晚期作为划分直立人与古老型智人的大致年代界限。近20年来,在非洲、欧洲和东亚新发现了一些更新世中期人类化石,目前古人类学界对中国更新世中期人类化石特征及演化有了与以往不同的认识。最近对大荔、许家窑、盘县大洞、许昌、华龙洞等人类化石的研究显示,近30万年以来东亚大陆人类演化呈现复杂的多样性,将这一时期人类全部归入古老型智人难以准确反映更新世中期中国古人类演化模式及规律。本文结合近年中国更新世中期人类演化研究进展,选择部分具有演化及分类价值的形态特征,分析这些特征在更新世中期中国古人类化石的表现特点。在此基础上,对更新世中期中国古人类演化模式做了尝试性探讨。本研究发现,周口店、和县、沂源、南京等中更新世早期人类化石呈现有较多的区域性特征,形态特征表现相对稳定;而大荔、金牛山、许家窑、许昌、华龙洞、马坝、盘县大洞等中更新世晚期人类在化石形态特征表现复杂多样,变异范围大。此外,在这一时期人类化石上发现较多与生存活动、健康、环境适应有关的证据。根据这些发现,作者认为中国中更新世早期组人类演化以形态连续性为主;进入中更新世晚期,中国古人类演化区域性特征减弱,演化模式以多样性为主。一系列新的化石发现和研究证据提示中更新世晚期东亚大陆可能生存有不同的古人类成员。根据目前掌握的化石形态和年代证据,大约30万年前是中国更新世中期演化变化关键时间节点。     

山西襄汾石沟砂场发现人类枕骨化石

2012年,山西襄汾县南贾镇上鲁村石沟砂场发现一件人类枕骨化石。该化石埋藏于汾河西岸第Ⅲ级阶地的砂砾层,地质时代可能是中更新世晚期。本文对枕骨形态特征进行描述,并将其与亚洲直立人、早期智人、早期现代人相关标本进行对比。观察发现,石沟古人类具有印加骨的形态特征,为中国古人类印加骨高频出现增添了新的证据;枕鳞中部没有发育枕骨圆枕,而具有最上项线,显示出枕骨形态的进步性,推测石沟古人类可能属于早期现代人。石沟古人类枕骨化石对于探讨中国及东亚地区古人类的系统演化问题提供了证据。     

Dental tissue proportions and enamel thickness in Neandertal and modern human molars

The thickness of dental enamel is often discussed in paleoanthropological literature, particularly with regard to differences in growth, health, and diet between Neandertals and modern humans. Paleoanthropologists employ enamel thickness in paleodietary and taxonomic studies regarding earlier hominins, but variation in enamel thickness within the genus Homo has not been thoroughly explored despite its potential to discriminate species and its relevance to studies of growth and development. Radiographic two-dimensional studies indicate that Neandertal molar enamel is thin relative to the thick enamel of modern humans, although such methods have limited accuracy. Here we show that, measured via accurate high-resolution microtomographic imaging, Neandertal molar enamel is absolutely and relatively thinner than modern human enamel at most molar positions. However, this difference relates to the ratio of coronal dentine volume to total crown volume, rather than the quantity of enamel per se. The absolute volume of Neandertal molar enamel is similar to that of modern humans, but Neandertal enamel is deposited over a larger volume of coronal dentine, resulting in lower average (and relative) enamel thickness values. Sample sizes do not permit rigorous intragroup comparisons, but Neandertal molar tissue proportions evince less variation than the modern human sample. Differences in three- and two-dimensional enamel thickness data describing Neandertal molars may be explained by dimensional reduction. Although molar tissue proportions distinguish Neanderthals from recent Homo sapiens, additional study is necessary to assess trends in tissue proportions in the genus Homo throughout the Pleistocene.     

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.