The Atapuerca sites and their contribution to the knowledge of human evolution in Europe

Over the last two decades, the Pleistocene sites of the Sierra de Atapuerca (Spain) have provided two extraordinary assemblages of hominin fossils that have helped refine the evolutionary story of the genus Homo in Europe. The TD6 level of the Gran Dolina site has yielded about one hundred remains belonging to a minimum of six individuals of the species Homo antecessor. These fossils, dated to the end of the Lower Pleistocene (800 kyr), provide the earliest evidence of hominin presence in Western Europe. The origin of these hominins is unknown, but they may represent a speciation event from Homo ergaster/Homo erectus. The TD6 fossils are characterized by a significant increase in cranial capacity as well as the appearance of a “sapiens” pattern of craniofacial architecture. At the Sima de los Huesos site, more than 4,000 human fossils belonging to a minimum of 28 individuals of a Middle Pleistocene population (ca. 500–400 kyr) have been recovered. These hominins document some of the oldest evidence of the European roots of Neanderthals deep in the Middle Pleistocene. Their origin would be the dispersal out of Africa of a hominin group carrying Mode 2 technologies to Europe. Comparative study of the TD6 and Sima de la Huesos hominins suggests a replacement model for the European Lower Pleistocene population of Europe or interbreeding between this population and the new African emigrants.     

Homo erectus—who,when and where: A survey

The state of information bearing on Homo erectus as developed since about 1960 is surveyed, with the resulting effects on problems. Definitions of H. erectus still rest on the Far Eastern samples (Chou-k'ou-tien/Java), and thus relate to late Lower to middle Middle Pleistocene material. Numerous important individual finds, however, have expanded the total: extension of the early and very early Sangiran material; very early to later in Africa, and relatively late in Europe. Datings remain uncertain or controversial within broad limits, but with some important successes and revisions. Discussion by authors of problems concerns degree of divergence among H. erectus populations and rate of evolutionary change; both appear relatively slight, but the data are inadequate for much present judgment. The apparent zone of transition to more advanced morphology (H. sapiens, sensu lato) by the late Middle Pleistocene better reflects signs of regional divergence. Some writers—not all—believe that even the earliest European fossils known (e.g., Petralona) had already advanced to a H. sapiens basic level, with later change in the direction of Neanderthals. A separate African phylum, from OH 9, is also suggested; recent Chinese finds may provide a third different post-erectus population before the Upper Pleistocene. Taxonomic expression of all this gives some problems.     

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.     

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。     

Homo erectus brain sizes by subspecies

Homo erectus fossils can be divided into four zoogeographic zones that show different rates of endocranial expansion during the Pleistocene. When these are also grouped into three time levels, we find small increases from early to middle forms, and regularly greater increases from middle to late forms. These increases fit a regular pattern that also accomodates all archaic types, including Neandertals, as late subspecies ofH. erectus.     

“Meganthropus” cranial fossils from Java

There are now twelve significant hominid cranial fossils from the Lower and Middle Pleistocene of Java, all but two being from the Sangiran site. Most of this material is well-known in the literature, but three skulls, possibly representing “Meganthropus” are here described in detail for the first time. Most scholars have assigned them all toHomo erectus, while others have suggested that they represent as many as four different hominoid taxa. The author argues that they represent two possible species of hominids. “Meganthropus” I, II, and III are more massive than any of the knownH. erectus specimens. They are also relatively higher vaulted, apparently smaller brained, and have unusually thick lower occipital planes. “Meganthropus” may represent a species that separated fromH. erectus upon its arrival to Java.     

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。     

A newHomo erectus cranium from Sangiran, Java

A newHomo erectus cranium was found on May 18, 1993 by Budi, a local farmer, at Sangiran. It dates from the Middle Pucangan Formation approximately 1.6–1.8 mya. The braincase is essentially complete and as is most of the face. The vault has the typicalH. erectus gable shape. There is a clear sagittal ridge beginning below the middle of the frontal squama and running to mid-parietal. Parasagittal ridges are rounded angulations halfway up the parietals, and coincide with poorly marked temporal lines. In all measurements, this skull is longer and consistently narrower than Trinil. It is chronologically and morphologically similar to the famousH. erectus skull from east Africa, KNMER-3733. Although existing much older, this new specimen is what one would expect a female counterpart to Sangiran 17 to look like.     

A SINGLE LINEAGE IN EARLY PLEISTOCENE HOMO: SIZE VARIATION CONTINUITY IN EARLY PLEISTOCENE HOMO CRANIA FROM EAST AFRICA AND GEORGIA

The relationship between Homo habilis and early African Homo erectus has been contentious because H. habilis was hypothesized to be an evolutionary stage between Australopithecus and H. erectus, more than a half‐century ago. Recent work re‐dating key African early Homo localities and the discovery of new fossils in East Africa and Georgia provide the opportunity for a productive re‐evaluation of this topic. Here, we test the hypothesis that the cranial sample from East Africa and Georgia represents a single evolutionary lineage of Homo spanning the approximately 1.9–1.5 Mya time period, consisting of specimens attributed to H. habilis and H. erectus. To address issues of small sample sizes in each time period, and uneven representation of cranial data, we developed a novel nonparametric randomization technique based on the variance in an index of pairwise difference from a broad set of fossil comparisons. We fail to reject the hypothesis of a single lineage this period by identifying a strong, time‐dependent pattern of variation throughout the sequence. These results suggest the need for a reappraisal of fossil evidence from other regions within this time period and highlight the critical nature of the Plio‐Pleistocene boundary for understanding the early evolution of the genus Homo.     

Three newHomo erectus mandibles from Java

There are now eleven manidublar pieces from the Lower and Middle Pleistocene of Java, all but one being from the Sangiran site. All of these have been assigned toHomo erectus by most workers, while others have suggested as many as four different hominoid taxa. Sangiran 21 (Mandible E), Sangiran 22 (Mandible F), and Sangiran 37 (Mandible G) are described here fully for the first time. Sangiran 21, 22, and 27 all come from the Upper Pucangan Formation and date approximately 1.2 Myr. The new mandibles are morphologically compatible with theH. erectus, crania from Java.     

The parietal bone of indonesian homo erectus

A comparative study of Indonesian parietal bones from Sangiran, Sambungmachan 1 and Ngandong has been undertaken. This study comprises a morphological and metrical analysis of the individual parietal bones, followed by consideration of the biparietal vault. The results are compared with other hominids from earlier and later periods. These hominids were found in China (Sinanthropus II, III, X, XI and XII), in Africa (ER 3733, OH 9, Ternifine, Broken Hill and Saldanha) and in Europe (Arago XLVII, Petralona, Swanscombe, Steinheim, Le Lazaret, La Chaise (Abri Suard) and Cova Negra). These European Middle Pleistocene hominids are attributed toHomo erectus by various authors (Lumley 1973;Hemmer 1972;Spitery 1982;Lumley andFournier 1982) and to an early Neanderthal group, pre-Neanderthal orHomo sapiens sensu lato (Neanderthals+modern humans) by others (Stringer 1980, 1981, 1983, 1984,Wolpoff 1980,Holloway 1982). The discussion about the classification of those hominids is not closed, but it is not the subject of this paper and not our intention to solve it here. So we have chosen to call this fossil material ‘Anteneandertals’ (Lumley 1973). It appears that some morphological metrical features allow us to separate the Sangiran and Ngandong samples. Sambungmachan 1, whose chronological age is not well established, appears to be closer to Ngandong men.     

Was Homo erectus responsible for the hand-axe culture?

This paper reviews the evidence from Africa, Asia and Europe of the cultural associations of Middle Pleistocene hominids, as well as the hominid skeletal associations of hand-axe remains.The author points out that it is possible to make a good argument—from the evidence of Steinheim, Kanjera and Swanscombe—that the hand-axes at these sites were made by Homo sapiens. On the other hand, on the basis of Fontéchevade and Vértesszöllös, it could be claimed that Middle Pleistocene Homo sapiens was responsible for primitive flake and chopper cultures. The evidence from Java is negative while that from China is directly opposed to the view that Homo erectus made hand-axes. Only from Ternifine in Algeria and Olduvai in Tanzania is there suggestive evidence that Homo erectus in those areas might have been responsible for the hand-axe culture. Thus, it is not possible at present to make any categorical statements as to the makers of either the great hand-axe culture or the flake and chopper culture, during Middle Pleistocene times.     

Cranial remains of Homo erectus from Beds II and IV,Olduvai Gorge,Tanzania

Cranial remains of hominids 9 and 12 from Olduvai Gorge are described in detail. O.H. 9 consists of a heavily built braincase, partly damaged and lacking the face, while O.H. 12 is less complete. The Bed II specimen is about 1.2 million years in age and shows anatomical similarities to the cranium designated ER-3733 from Koobi Fora, east of Lake Turkana. Together these African fossils provide valuable information about Homo erectus in the later Lower Pleistocene. Comparisons of O.H. 9 with several of the Choukoutien crania are also carried out. These Chinese and other Asian remains of Homo erectus cannot be placed in a secure chronological framework, but all of the material should be studied systematically in order to assess relatedness among what must be several different populations.     

Reconstructing cranial evolution in an extinct hominin

Homo erectus is the first hominin species with a truly cosmopolitan distribution and resembles recent humans in its broad spatial distribution. The microevolutionary events associated with dispersal and local adaptation may have produced similar population structure in both species. Understanding the evolutionary population dynamics of H. erectus has larger implications for the emergence of later Homo lineages in the Middle Pleistocene. Quantitative genetics models provide a means of interrogating aspects of long-standing H. erectus population history narratives. For the current study, cranial fossils were sorted into six major palaeodemes from sites across Africa and Asia spanning 1.8–0.1 Ma. Three-dimensional shape data from the occipital and frontal bones were used to compare intraspecific variation and test evolutionary hypotheses. Results indicate that H. erectus had higher individual and group variation than Homo sapiens, probably reflecting different levels of genetic diversity and population history in these spatially disperse species. This study also revealed distinct evolutionary histories for frontal and occipital bone shape in H. erectus, with a larger role for natural selection in the former. One scenario consistent with these findings is climate-driven facial adaptation in H. erectus, which is reflected in the frontal bone through integration with the orbits.     

Frontal bone fragment of<Emphasis Type="Italic">Homo erectus</Emphasis> from Sangiran,Java

In 1994 a hominid frontal bone fragment was found in the river floor of the Brangkal River, the Sangiran area, Central Java. The original stratigraphic level is not known at present stage of the research. But it is possible that the bone was derived from the Grenzbank zone of the Bapang Formation (Lower/Middle Pleistocene). Morphological features of the bone, such as a thick and continuous supraorbital torus, a wide and flat supratoral plane, and a flat and strongly inclined frontal squame suggest that the bone is assigned to JavaneseHomo erectus, especially to the Sangiran and Trinil group of it.     

Morphological and functional aspects of the temporal bone articular tubercle morphology inHomo erectus andHomo sapiens

The morphological variability of the temporal articular tubercle was studied inHomo erectus andHomo sapiens. Five configurations have been defined. There is a high heterogeneity amongHomo erectus. The Neandertal lineage and that leading toHomo sapiens sapiens are more homogenous, each of them exhibits a high frequency of one configuration. This study has also focused on the functional implications of this variation. A theoretical approach to two different configurations of the articular tubercle is considered in the same bony, muscular and ligamentary context. This suggests that the configuration which consists of a transverse concavity is, for the mandible depression, concomitant with a slight functional disadvantage in comparison with the cylindrical configuration. It appears that a midfacial projection allows for a compensation of this disadvantage. It is concluded that this model can be proposed for Neandertals which present a very concave articular tubercle and a typical midfacial projection.