High-precision U-series dating of Locality 1 at Zhoukoudian, China

Thermal ionization mass spectrometric(230)Th/(234)U dating has been carried out on intercalated speleothem samples from the limestone cave occupied by Homo erectus at Zhoukoudian, China. The samples were recently collected in proper stratigraphic context after detailed field examinations. The results show that the age of the No. 5 Skull from Layer 3 is >400 ka, possibly in the range of about 400-500 ka, and that the hominid fossils from the lower strata are at least 600 ka and possibly >800 ka, much older than previously thought. The near-equilibrium(230)Th/(234)U ratios and internal consistency of the dates and stratigraphy lend credence to the results and allow us to comment on their important implications for human evolution.     

印度纳尔玛达地区发现的粗壮智人化石及其与亚洲直立人的连续性

1999年,在印度西部的纳尔玛达(Narmada)河的一条支流——奥尔桑(Orsang)河谷发现了一具孤立的智人头骨,这对于了解印度人类进化历史和南亚直立人与最古老的亚洲智人(或AMH,解剖学上现代的人)之间的联系有着重要的意义。化石是在古老的河流沉积物中发现的。对主体沉积和颅内沉积物的红外光释光(IRSL)测年结果显示,其年代为3—5万年。然而,对化石的直接测年(放射性碳加速器质谱测年AMS)结果表明,其年代最小为4981—5579年前。头骨被归类为圆颅型智人。头骨上最引人注意的特征是颅外有亚洲直立人头后部很发达的角圆枕。头骨最宽处在下部(颞骨)如直立人,这样的情况从来不见于智人。这可能是由于乳突的气窦化而形成的。乳突上脊发育,从破损的眉间区(20mm)可以看到很宽的额窦。所有这些粗壮的特征都表明Orsang头骨和晚期亚洲直立人之间具有遗传连续性。     

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.     

The subspecies ofHomo erectus

ClassifyingHomo erectus into subspecies can be based on either temporal or geographical differences, but there is no accepted system for using both. This can be done with subspecies names consisting of two elements — a prefix ofneo, meso, orpaleo to indicate grade, followed by a geographical term ofeuropus, africus, sinicus, orindicus to indicate line. Thus Rhodesian isHomo erectus neoafricus, Ngandong isHomo erectus neoindicus, Peking isHomo erectus mesosinicus, ER 3733 isHomo erectus paleoafricus, etc.     

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.     

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.     

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。     

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 face of Olduvai Hominid 12

Facial remains of Homo erectus are rare and their scarcity hinders our understanding of the variability and relationships in this taxon. Previously undescribed fragments of the peri-orbital region and unidentified matches between fragments of Olduvai Hominid 12 (OH 12) enhance comparison of the African H. erectus hypodigm. The newly reconstructed upper face and maxilla of OH 12 is most similar in size and shape to that of KNM-ER 3733, despite being as much as one million years younger than the Koobi Fora hominin. However, the posterior vault and mastoid region of OH 12 are most similar to OH 9. This combination of morphology suggests that the relationship between the Olduvai and Koobi Fora portions of the H. erectus hypodigm requires reconsideration.     

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。     

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。     

Poor fossil record and major changes around 1 MaBP

The abundance of early fossil humans in African sites ceases at dates around 1.3MaBP; there is almost none until nearly 0.8MaBP. Again these are scarce until less than 0.5 Ma. Most of Homo erectus fossils in Java are dated between c.1.3 and 0.70Ma; just a few fossil humans are known in Eurasia for this time span. Questions arise on eventual environmental constraints, that may have influenced evolutionary crises of human populations, but also on geographic conditions adverse to fossilization processes and/or site formation. Records on climate variables, vegetation, sea level, sedimentary conditions and tectonic behavior in regions of the Old World are collected, and correlation traced back in time slices. Continental accretion is related to end of sedimentation in African basins; repeated compressive tectonic activity negatively affected preservation of both fossils and occupation sites widely, before and immediately after the M/B reversal. Lithic artefact assemblages, although derived, are understood as evidence of human occupation in the studied interval. Human populations are exspected to be affected by environmental changes, small, and scattered. Origin of Homo sapeins and divergence of preneandertalians are driven back to those times. Ancestorship to the latter is found probably in African fossils of that time span. Modern traits found in the Atapuerca-TD6 sample and the preceeding considerations urge research on humans and their movements intra- and intercontinental around 1 MaBP.     

Cladistic analysis of early Homo crania from Swartkrans and Sterkfontein, South Africa

The phylogenetic relationships of early Pleistocene Homo crania from the South African sites of Swartkrans and Sterkfontein were investigated through cladistic analyses of 99 morphological characters. The Swartkrans Member 1 specimen SK 847 and the Stw 53 cranium from Sterkfontein Member 5A were treated as separate operational taxonomic units (OTUs), distinct from the three species of early Homo-H. erectus, H. habilis, and H. rudolfensis-that are recognized from the Plio-Pleistocene deposits of East Africa. The cladistic analyses differed in the treatment of the South African OTUs (separate Swartkrans and Sterkfontein OTUs vs. a single Swartkrans+Sterkfontein OTU). PAUP 4.0 was used to construct cladograms and address hypotheses about relationships. In the analysis that treated the South African specimens as a single OTU, the position of that OTU was stable as a separate branch on the Homo clade between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)]. When SK 847 and Stw 53 were treated as separate OTUs, the majority of most parsimonious trees indicated that they were positioned in similar positions as the combined South African Homo OTU; that is, as separate branches between H. rudolfensis and [H. habilis+(H. erectus+H. sapiens)], with the Swartkrans OTU generally occupying a more derived position. The position of the Sterkfontein OTU was more stable than that of the Swartkrans OTU, which was found in several other positions among the minimum length trees. Running the analyses with only those characters preserved by SK 847 and Stw 53 resulted in similar topologies for minimum length trees, although the positions of Stw 53, SK 847, and H. habilis exchanged places in some trees. In no case was an exclusive sister relationship between either South African OTU and a particular species of Homo supported statistically. Both South African OTUs differ from H. habilis in the fewest number of cladistic characters.     

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,     

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.     

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。     

南京汤山直立人颅容量的推算

以耳上颅高推算现代人颅容量的公式可适用于汤山直立人1号头骨的颅容量的推算。汤山直立人1号头骨的颅容量推算值为871ml,与周口店直立人3号头骨的颅容量值相近,提示了汤山直立人的年代“早于40万年或更早”的看法更可取些。