古蛋白质分析在东亚古人类演化中的应用前景

东亚古人类演化是学术界关注的热点科学问题,国内外学者对此进行了多学科的相关研究,取得了很多重要进展,但仍然存在许多尚未解决的问题。古蛋白质分析近年来成为古生物演化领域的又一个前沿和热点方向,取得了一系列重要突破。较之古DNA,古蛋白质的保存优势使其可以在时间上和地域上突破古DNA的限制,在古人类演化领域大有可为。东亚古人类化石丰富且时段大致连续,但更新世或更早时期的分子证据非常缺乏。本文从古蛋白质分析的发展史、研究潜力、难点与挑战以及思考与展望等几方面,对古蛋白质分析在东亚古人类演化研究中的应用前景进行梳理与思考。相信随着更多分子证据的积累,古蛋白质分析可为东亚古人类的演化脉络提供更多关键性的线索,极大地促进人类演化研究。     

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。     

泥河湾盆地早更新世山神庙咀遗址动物群及其时代意义

狭义(或经典)泥河湾动物群主要产自桑干河北岸的下沙沟村周边,化石来源于不同地点;该动物群被作为我国北方早更新世的代表,但其绝对年代一直饱受争议,最新古地磁测年数据是2.2-1.7 MaBP。由于受河流及断层切割影响,泥河湾盆地内部不同区域尤其是桑干河两岸的地层对比,一直是个科学难题,其根本原因在于长期以来在桑干河南岸未发现重要化石点。最近,在桑干河南岸的岑家湾台地发现了山神庙咀化石点,目前已在此地点做过9次正规发掘,出土哺乳动物化石1526件,鉴定出25个种(含未定种),其中23种属于狭义泥河湾动物群的成员。由此看出,山神庙咀动物群的时代应当与狭义泥河湾动物群的时代接近,其地质年代应当不晚于1.7 MaBP。山神庙咀遗址与小长梁遗址隔沟相望,但其化石层稍高于小长梁文化层;山神庙咀动物群的研究结果还表明,小长梁遗址年代要比先前所测得的1.36 MaBP要早;小长梁周边的史前遗址的地质年代应当再做深入研究。小长梁地区的泥河湾层底部普遍存在砾石层,但其出现的水平位置以及砾石在分选和磨圆程度等方面都各不相同。调查清楚不同剖面中底砾层的分布状况及其性质,将会对本地区地层对比研究以及推断当时泥河湾...     

Middle pleistocene humans from africa

Patterns of human evolution in the Middle Pleistocene remain poorly understood. There is general consensus that by the onset of this time period, populations ofHomo erectus were dispersed from Africa into Eurasia, including the Far East. In the western part of this range (perhaps in Africa),Homo erectus then produced a daughter lineage exhibiting more advanced characters of the face, braincase and cranial base. How this new species should be defined is currently debated. In my view, fossils from sites such as Bodo and Broken Hill in Africa may be lumped with material from earlier Middle Pleistocene localities in Europe. Such a taxon is appropriately namedHomo heidelbergensis. Whether the hypodigm should be extended to include fossils from China is another question. In any case, this group of hominids is plausibly ancestral to both the specialized Neanderthals of Europe and more modern humans of the later Middle Pleistocene.     

A new pleistocene hominid-bearing locality at Hoedjiespunt,South Africa

HDP1 is an archaeological and faunal site located on the Hoedjiespunt peninsula at Saldanha Bay, South Africa, that has recently yielded fossil human remains. Artefacts from the associated archaeological deposits are identified as being Middle Stone Age. U series analysis of capping calcretes and analysis of the foraminifera and fauna associated with the human fossils indicate an age for the deposit in excess of 74,000 years before present, and it most probably dates to around 300,000 years before present. The fossil human teeth from in situ deposits at Hoedjiespunt are described and found to be large by comparison with modern humans but smaller than the known upper dentitions of southern African “archaic” Homo sapiens. The Hoedjiespunt molars are found to be morphologically within the range of variation observed in the teeth of modern Homo sapiens. © 1995 Wiley-Liss, Inc.     

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。     

Human evolution

The common ancestor of modern humans and the great apes is estimated to have lived between 5 and 8 Myrs ago, but the earliest evidence in the human, or hominid, fossil record is Ardipithecus ramidus, from a 4.5 Myr Ethiopian site. This genus was succeeded by Australopithecus, within which four species are presently recognised. All combine a relatively primitive postcranial skeleton, a dentition with expanded chewing teeth and a small brain. The most primitive species in our own genus, Homo habilis and Homo rudolfensis, are little advanced over the australopithecines and with hindsight their inclusion in Homo may not be appropriate. The first species to share a substantial number of features with later Homo is Homo ergaster, or ‘early African Homo erectus’, which appears in the fossil record around 2.0 Myr. Outside Africa, fossil hominids appear as Homo erectus-like hominids, in mainland Asia and in Indonesia close to 2 Myr ago; the earliest good evidence of ‘archaic Homo’ in Europe is dated at between 600–700 Kyr before the present. Anatomically modern human, or Homo sapiens, fossils are seen first in the fossil record in Africa around 150 Kyr ago. Taken together with molecular evidence on the extent of DNA variation, this suggests that the transition from ‘archiac’ to ‘modern’ Homo may have taken place in Africa.     

The current state of korean paleoanthropology

The hominid fossil and Paleolithic archaeology records from the Korean Peninsula are extensive, but relatively little is known about the Korean human evolutionary record outside this region. The Korean paleoanthropological record is reviewed here in light of major research issues, including the hominid fossil record, relative and chronometric dating, lithic analysis, hominid subsistence, and the presence of bone tools, art and symbolism. Some of the major conclusions drawn from this review include: (1) hominid fossils have been found in nine separate sites on the Korean Peninsula; (2) possible Homo erectus fossils are present in North Korea; (3) Ryonggok Cave, in North Korea, has exposed the remains of at least five archaic Homo sapiens individuals; (4) a possible burial of an anatomically modern Homo sapiens child, discovered in Hungsu Cave in South Korea, has been tentatively dated to roughly 40,000 years ago; (5) handaxes and cleavers have been found at a number of sites near Chongokni and they appear to date to at least 100,000 years ago; and (6) taphonomic studies are necessary for addressing issues related to determining the nature of hominid-carnivore interaction over similar resources (e.g. carcasses and shelter); and the presence/absence of Early Paleolithic bone tools, art, and symbolism in Korea.     

New hominid fossils from the Swartkrans formation (1979-1986 excavations): postcranial specimens

New postcranial fossils of Paranthropus robustus and Homo cf. erectus were recovered from Swartkrans from 1979 through 1986. These fossils are from Members 1, 2, and 3. The new fossils are described here along with their morphological affinities. Fossils that are assigned to Paranthropus indicate that the South African "robust" australopithecines engaged in tool behavior and were essentially terrestrial bipeds at around 1.8 Myr BP. The manual dexterity and bipedal locomotion of Paranthropus may have equaled that of Homo habilis in East Africa at approximately the same time.     

Middle and late Pleistocene faunas of Pinnacle Point and their paleoecological implications

The Western Cape region of South Africa is home to a unique type of mediterranean vegetation called fynbos, as well as some of the earliest sites of modern human occupation in southern Africa. Reconstructing the paleohabitats during occupations of these early anatomically modern Homo sapiens is important for understanding the availability of resources to the humans during the development of behaviors that are often considered advanced. These reconstructions are critical to understanding the nature of the changes in the environment and resources over time. Here we analyze the craniodental fossils of the larger mammals recovered from two Pleistocene assemblages in the Pinnacle Point complex, Mossel Bay, Western Cape Region, South Africa. We reconstruct the paleohabitats as revealed by multivariate analyses of the mammalian community structures. Pinnacle Point 30 is a carnivore assemblage and Pinnacle Point 13B includes early evidence of a suite of modern human behavior; together they present an opportunity to identify environmental change over time at a localized geographic scale. Further, this is the first such study to include dated Western Cape localities from Marine Isotope Stage 6, a time of environmental pressure that may have marginalized human populations. Results indicate that environmental change in the Western Cape was more complex than generalized C(4) grassland expansions replacing fynbos habitats during glacial lowered sea levels, and thus, resources available to early modern humans in the region may not have been entirely predictable.     

The Atapuerca human fossils

After 20 years of research, the Atapuerca sites have provided a large amount of archaeological and palaeontological remains. Human fossils have been found in three sites: Gran Dolina, galería and Sima de los Huesos. The Early Pleistocene human fossils from Gran Dolina have been ascribed to a new species,Homo antecessor, that represent the last common ancestor of Neandertals and modern humans. The Sima de los Huesos fossils and all the European Middle Pleistocene human fossils are the ancestors exclusively of the Neandertals, which evolved in Europe in conditions of geographic and genetic isolation.     

Metric and non-metric randomization methods, geographic variation, and the single-species hypothesis for Asian and African Homo erectus

This paper proposes a statistical test of the single-species hypothesis using non-metric characters as a complement to statistical tests using more traditional metric characters. The sample examined is that of Asian and African Homo erectus. The paleoanthropological community is divided on the taxonomic distinction of these fossils, with workers arguing both for and against the species-level distinction between Asian and African populations. Previous arguments have focused on patterns of apparent morphological differentiation between the African and Asian cranial samples. To assess this question, three tests were performed that compared the range of variation in the fossil sample to a single-species group with a similar geographic distribution; this comparative sample was composed of 221 modern humans from Africa and Asia. For the first test, 23 metric characters were analyzed on the fossil and comparative samples. Using resampling procedures, the variation for these characters was examined, recreating 1000 samples from the human analogs and comparing the CV distributions of these samples to the CVs of the fossil group. The second test used the metric data to calculate a Euclidean distance between the African and Asian fossil samples. This distance was compared to a distribution of Euclidean distances calculated between 1000 randomly selected samples of African and Asian modern humans. For the third test, a grading scale was created for ten non-metric characters that encompassed the total morphological variation found in the fossil and modern human samples. The Manhattan distance between the Asian and African fossil samples was calculated and compared to a distribution of distances calculated between 1000 randomly selected samples of African and Asian moderns. The first two tests, using the metric data, failed to falsify the null hypothesis. However, in the third test, using non-metric data, the total Manhattan distance for the fossil sample approached the 100th percentile of the resampled distances calculated from the moderns. The implications of the contrasting results are discussed.     

African ape ancestry

It is commonly believed that the australopithecines are more closely related to humans than to African apes. This view is hardly compatible with the biomolecular data which place theHomo/Pan split at the beginning of the australopithecine period. Nothing in the fossil hominid morphology precludes the possibility that some australopithecines were ancestral to gorillas or chimpanzees and others to humans.     

Middle eastern origin model forHomo sapiens (Moderns & Neanderthals), language and modern behaviour

A new model may resolve the problem of when and where did appear anatomically modern humans. According to this model, Neanderthals were probably neither our ancestor nor different species.Homo sapiens appeared probably in the Middle East, approximately 150 ka ago and differentiated to anatomically modern humans and Neanderthals because of the genetic programme. The fossils older than 150 ka are probably not Neanderthal such as Zuttiyeh and Biache-Saint-Vaast specimens. Cultural capacities of Neanderthals were probably equivalent to Moderns. Most of pre-Homo sapiens populations may be extinct without replacement byHomo sapiens. Language and modern behaviour should have arisen with our own species.     

20世纪的中国人类古生物学研究与展望

本文回顾中国人类古生物学在20世纪的主要研究进展。中国的化石曾经数度入围最早期的人类。目前中国已有60余处地点发现了人类化石, 其中最古老的暂时要推云南元谋上那蚌的两颗门齿。对中国人类化石的研究提出了连续进化附带杂交的模式, 支持现代人起源的多地区进化说。中国各时期的化石人类有较显著的地区差异。中国人类的发展可能比喻为既有分支间的杂交融合, 又有小分支的绝灭和局部替代的河网状。     

The human cranial remains from Gran Dolina Lower Pleistocene site (Sierra de Atapuerca, Spain).

In this article we study the cranial remains of the late Lower Pleistocene human fossils from Gran Dolina (Sierra de Atapuerca, Spain), assigned to the new species Homo antecessor. The cranial remains belong to at least five individuals, both juveniles and adults. The most outstanding feature is the totally modern human morphology of the very complete face ATD6-69, representing the earliest occurrence of the modern face in the fossil record. The Gran Dolina fossils show in the face a suite of modern human apomorphies not found in earlier hominids nor in contemporary or earlier Homo erectus fossils. There are also traits in the Gran Dolina fossils shared with both Neandertals and modern humans, which reinforce the hypothesis that Neandertals and modern humans form a clade, and that the Gran Dolina fossils are a common ancestor to both lineages.     

Did a discrete event 200,000-100,000 years ago produce modern humans?

Scenarios for modern human origins are often predicated on the assumption that modern humans arose 200,000-100,000 years ago in Africa. This assumption implies that something ‘special’ happened at this point in time in Africa, such as the speciation that produced Homo sapiens, a severe bottleneck in human population size, or a combination of the two. The common thread is that after the divergence of the modern human and Neandertal evolutionary lineages ∼400,000 years ago, there was another discrete event near in time to the Middle-Late Pleistocene boundary that produced modern humans. Alternatively, modern human origins could have been a lengthy process that lasted from the divergence of the modern human and Neandertal evolutionary lineages to the expansion of modern humans out of Africa, and nothing out of the ordinary happened 200,000-100,000 years ago in Africa.Three pieces of biological (fossil morphology and DNA sequences) evidence are typically cited in support of discrete event models. First, living human mitochondrial DNA haplotypes coalesce ∼200,000 years ago. Second, fossil specimens that are usually classified as ‘anatomically modern’ seem to appear shortly afterward in the African fossil record. Third, it is argued that these anatomically modern fossils are morphologically quite different from the fossils that preceded them.Here I use theory from population and quantitative genetics to show that lengthy process models are also consistent with current biological evidence. That this class of models is a viable option has implications for how modern human origins is conceptualized.     

A fossil hominoid proximal femur from Kikorongo Crater, southwestern Uganda

The external morphology of a fragmentary right proximal femur from southwestern Uganda is described here. Discovered in the Kikorongo Crater of Queen Elizabeth National Park in 1961, this specimen was informally assigned to Homo sapiens (although never described) and tentatively dated to the late Pleistocene. However, because aspects of the external morphology of the femur align the fossil with the African great apes, we suggest that the Kikorongo femur may be the first postcranial fossil of the genus Pan. Like the African apes, the Kikorongo specimen lacks both an obturator externus groove and an intertrochanteric line. It has a short femoral neck with a circular cross section, and a narrow and deep superior notch. Using resampling statistics and discriminant function analysis, the Kikorongo femur clustered with the genus Pan, as opposed to Gorilla or Homo. However, if the specimen is from Pan, it would be large for this taxon. Furthermore, features that clearly distinguish the external morphology of Plio-Pleistocene hominin proximal femora from African ape femora, such as the shape of the femoral neck in cross section and femoral neck length, have converged in Holocene humans and African apes. Unfortunately, the internal morphology of the femoral neck of the Kikorongo fossil was not discernable. Although we hypothesize that the Kikorongo femur is from the genus Pan, there is such variability in the proximal femora of modern humans that, although it would be an unusual human, it remains possible that this fossil represents H. sapiens.     

广西柳江土博甘前洞的铀系年代

本文报道晚期智人化石地点广西柳江博甘前洞新生碳酸盐岩和骨化石样的铀系测年结果。该地点表层钙板在约94Ka前开始形成,含化石粘土堆积叠压的钙板年代为约20ka,人牙化石应位于二者之间。二个动物化石样的二种铀系法年代范围为85－139ka,表明该地点与含化石堆积与表层钙板间无地层倒序现象,支持人牙化石大于100ka的结论。邻近的柳江晚期智人化石地点和柳州白莲洞人类遗址铀系测年的结果与本文一致。具现代解剖特征智人在中国南方出现的时间,很可能不晚于西非和非洲。在现代人类起源方面,中国不应是远离中心、滞后和被取代的地区。