Nasal morphology and the emergence of Homo erectus

Modern humans, among extant hominoids, possess a unique projecting, external nose whose basic structure is reflected in a series of skeletal features including nasal bone convexity, an internasal angle, lateral nasal aperture eversion, prominence and anterior positioning of the anterior nasal spine, an acute angle of the subnasal alveolar clivus, and an expansion of the breadth of the nasal bones relative to that of the piriform aperture. This anatomy appears with the emergence of Homo erectus ca. 1.6 million years ago. Although it undoubtedly evolved in the context of craniofacial and dental reduction during hominid evolution, it appears to have been primarily a response to the need for moisture conservation in an arid environment via turbulence enhancement and ambient cooling of expired air. Its appearance at this time in hominid evolution, in conjunction with the presence of a fatigue-resistant locomotor anatomy characteristic of archaic members of the genus Homo, indicates a shift to increasingly prolonged bouts of activity in open and arid environments.     

周口店直立人3号头骨的性别和年龄:南京直立人头骨的启示

周口店直立人3号头骨是1929年发现的, 被步达生鉴定为一青春期或成年期的女性个体的头骨。后来, 步达生改变了观点, 认为该头骨是代表青春期早期的男性个体。魏敦瑞同意步达生的意见, 但认为该头骨应属于8岁或9岁的男性个体。此后,周口店直立人3号头骨的性别和年龄的鉴定一直困扰着许多古人类学家。1993年发现的南京直立人成年女性头骨在形态上和尺寸上与周口店直立人3号头骨有相近之处。本文对周口店直立人3号头骨与南京直立人等头骨作了形态和测量上的比较, 结果表明: 从头骨的尺寸大小, 眶上圆枕、枕圆枕和肌嵴的发育程度以及头骨骨壁厚度等来看, 周口店3号头骨有理由被认为是女性个体的。该头骨的鼓骨裂、下颌骨关节窝、泪腺窝、额窦、骨缝等都无法证明其是幼年个体。周口店直立人3号头骨应代表成年女性个体。     

A comparison of the nariokotome Homo erectus with juveniles from a modern human population.

The Nariokotome Homo erectus has an apparent disjunction of inferred age as judged by dental maturity, by epiphyseal closure and by stature, when compared to modern human norms. On this basis, it has been suggested that this fossil hominin differed in its pattern of growth and development from modern humans. In particular, the characteristic human adolescent growth spurt may not yet have been present, and in this sense H. erectus growth would be more ape-like than human-like. This study tests this conclusion by examining the variation in age as inferred from the maturity indicators in a modern human skeletal population of known age. The results show that all of the maturity indicators used in this analysis underage the test skeletons. Furthermore, there is also no consistency between the indicators; they do not agree in their inferred chronological ages. The disjunction between the maturity indicators in the test skeletons is similar in pattern to the disjunction observed in the Nariokotome Homo erectus. This is particularly true of the relationship between dental age and the other two indicators. These results suggest that the pattern observed in Nariokotome is within the normal range of variation found in modern humans. It does not necessarily indicate a different pattern of growth and development.     

A re-evaluation of the metric diversity within Homo erectus

Previous work by several researchers has suggested that the cranial sample from Zhoukoudian possesses a unique metric pattern relative to the African and Asian specimens assigned to Homo erectus. The current study readdresses this issue with an expanded fossil sample and a larger and more comprehensive set of cranial measurements. To test the patterns present in the assemblage, canonical variates analysis was performed using a covariance matrix generated from the Howells data set. From this, interindividual Mahalanobis distances were computed for the fossils. Random expectation statistics were then used to measure statistical significance of the Mahalanobis distances. The results show that the Zhoukoudian hominids exhibit a unique metric pattern not shared by the African and Indonesian crania sampled. In these tests the Hexian calvaria resembled the African and Indonesian specimens and differed significantly from the craniometric pattern seen in the Zhoukoudian fossils. The Zhoukoudian specimens are characterized by a wide midvault and relatively narrow occipital and frontal bones, while the African and Indonesian crania (including Hexian) have relatively broad frontal and occipital dimensions compared to their midvaults. These results do not suggest that a multiple-species scenario is necessary to encompass the variation present in the sample. Based on the current evidence it is more probable that this variation reflects polytypism influenced by environmental adaptation and/or genetic drift.     

Fractal dimension of the middle meningeal vessels: variation and evolution in Homo erectus, Neanderthals, and modern humans

The middle meningeal vascular network leaves its traces on the endocranial surface because of the tight relationship between neurocranial development and brain growth. Analysing the endocast of fossil specimens, it is therefore possible to describe the morphology of these structures, leading inferences on the cerebral physiology and metabolism in extinct human groups. In this paper, general features of the meningeal vascular traces are described for specimens included in the Homo erectus, Homo neanderthalensis, and Homo sapiens hypodigms. The complexity of the arterial network is quantified by its fractal dimension, calculated through the box-counting method. Modern humans show significant differences from the other two taxa because of the anterior vascular dominance and the larger fractal dimension. Neither the fractal dimension nor the anterior development are merely associated with cranial size increase. Considering the differences between Neanderthals and modern humans, these results may be interpreted in terms of phylogeny, cerebral functions, or cranial structural network.     

南京直立人的高耸鼻梁和气候适应

为论证南京直立人高耸的鼻梁是对气候的适应,本文检验了有关鼻骨的3种测量项目;结果表明:鼻尖指数和鼻根指数用来表达整个骨性鼻梁的外突程度是有价值的,但眶间宽高指数不能用来说明整个鼻梁外突程度。Carey和Steegmann氏著作中的各人群鼻尖指数和鼻根指数数据和相应的气候资料表明,干燥地区人群的鼻比潮湿地区的要较外突,寒冷地区人群的鼻要比温暖地区的较外突。这种鼻的外突程度与干、寒气候的关系可以从人鼻的解剖学和呼吸生理学方面得到支持。因纽特人(爱斯基摩人)虽然鼻梁并未达到高耸级别,但有因上颌窦缩小而明显扩大的鼻腔,大的鼻甲、鼻道等。因此,因纽特人的鼻形态代表鼻对寒冷气候适应的另一种方式,不能用作证据来否认鼻梁高耸与气候之间的关系。另外,对南京直立人化石地点的孢子花粉、植硅体的研究表明,南京直立人是生活在冰期环境中,当时的冰期在很大程度上可以与冰期盛期或大冰期相对比。综上所述,南京直立人鼻梁高耸是对寒冷气候适应的结果,未必是由于基因交流的结果;这是对南京直立人鼻梁高耸的成因的目前一个最合理的解释。     

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。     

Axial and appendicular skeleton of Homo antecessor.

The human trunk and limb bones recovered from the Gran Dolina site, in the Sierra de Atapuerca (Burgos, Spain) are studied. All these fossils were excavated at the level called TD6 between 1994 and 1995 and have been dated in excess of 780,000 years ago. These remains have been recently attributed to a new Homo species named Homo antecessor. Axial (vertebrae and ribs) and part of the appendicular (clavicles, radii, femur and patellae) skeleton are studied here. Hand and foot bones have been studied elsewhere (Lorenzo et al., 1999). Four is the minimum number of individuals represented by the postcranial remains recovered up to now. All elements are briefly described anatomically, measured and compared with other fossil hominids and modern humans in order to establish, as far as possible, what postcranial morphology characterized this new species of our genus.The H. antecessor postcrania, generally, display a set of morphological traits that are more similar to modern humans than to the Middle and Upper Pleistocene European hominids. Our results do not contradict the previous phylogenetic analysis, i.e., that H. antecessor represents the last common ancestor for H. sapiens (modern humans) and H. neanderthalensis (Neandertals).     

再论南京直立人的高耸鼻梁和气候适应

对南京直立人所具有的高耸鼻梁, 本文作者曾从人鼻的功能、鼻梁高耸程度与气候因素的关系以及因纽特人的鼻对寒冷气候的适应等方面进行过论证,认为是对气候适应的结果。本文从南京直立人的生存年代、若干形态项目的可靠性、人类的适应性以及高耸鼻梁的化石人类谱系对这一问题作进一步的论证。结果表明: 1)尚无可靠的证据能够否定南京直立人是生活在距今58—62万年这一年代测定数据。在南京直立人生存时,欧洲远古人类尚未具有像南京直立人那样的高耸鼻梁; 2)人类高耸鼻梁这一适应性特征的形成会有永久性的影响, 是遗传特征, 其后代即使生活在与其祖先不同的气候环境里, 这个特征也不会马上改变; 3)如果有所谓的Bodo高鼻梁的化石人类谱系存在的话, 则南京直立人的高耸鼻梁与欧洲化石人类的基因或与这个Bodo谱系的基因并无遗传上的关系。一个合理的解释是, 南京直立人的高耸鼻梁不可能意味着西方血统。     

周口店直立人头骨创伤与人工切割痕迹辨析

自20世纪40年代魏敦瑞提出北京猿人（周口店直立人）可能存在“暴力争斗”和“同类残食”的观点之后,引发了一些学者和科普大众对北京猿人是否为“食人族”的争论。由于原始化石标本在第二次世界大战期间丢失,本文以保存下来的周口店（ZKD）直立人头骨原始模型和素描图为研究材料,对魏敦瑞提出的具有创伤和人工切割痕迹的5件标本（ZKD II、ZKD VI、ZKD X、ZKD XI和ZKD XII）进行辨析。结果显示：1）ZKD X、ZKD XI、ZKD XII这3件标本的头盖部,有7处可以确定为生前遭受非致死性撞击导致的局部创伤;2）ZKD VI头盖骨残片周边的及中央区域的断痕,为个体生前遭受致死性的暴力打击导致;3）ZKD II和ZKD VI顶骨位置疑似人工切割痕迹的沟槽实际上是动物啃咬或自然因素导致的,北京猿人“同类残食”的观点在本文研究中没有得到证实。北京猿人头骨的创伤痕迹都位于头盖部,以顶骨居多,其次为额骨,符合人群之间暴力冲突产生创伤的位置。     

On the paleopathologic findings exhibited by the late Homo erectus of Ceprano, Italy

In March 1994, during excavation work for the construction of a motorway, a fragmented calvaria was discovered in the “Campo Grande” area near Ceprano, a town in southern Latium (Central Italy) about 55 miles from Rome. After reconstruction, the remain was recognized as belonging to aHomo erectus; it has been estimated that it goes back to the lowest middle Pleistocene. The calvaria exhibits two pathologic findings. The first is a congenital malformation of the sphenoidal sinus consisting in a deep, wide recess penetrating into the left greater wing as far as the sphenotemporal suture. The lesion was examined with the help of a cast whose features were compared with those described in some early and recent cases reported in the literature. The second finding is a healed, depressed fracture of the right brow ridge. The cause and mechanism of this lesion are discussed, and a mechanical approach has been used to provide information pertinent to the specific reason for the injury. The possibility that the hominid was butted by a large animal appears to be the most likely cause of the fracture.     

Energetics in Homo erectus and other early hominins: the consequences of increased lower-limb length

Previous studies of daily energy expenditure (DEE) in hominin fossils have estimated locomotor costs using a formula that was based on six species, all 18 kg or less in mass, including no primates, and that has a number of other problems when applied in an ecological context. It is well established that the energetic cost of human walking is lower than that of representative mammals, particularly for individuals with long lower limbs. The current study reevaluates the daily energy expenditures of a variety of hominin species using more appropriate approaches to estimating locomotor costs. To estimate DEE for primates, I relied on published data on body mass, day range, and the percentage of time spent in various activities. Based on those data, I calculated a value for nonlocomotor DEE. I then used a variant of a method that I have suggested elsewhere to calculate the daily cost due to locomotion (DEEL) and summed the two to calculate total DEE. The more up-to-date methods for calculating the cost of travel result in lower estimates of this aspect of the energy budget than seen in previous studies. Values obtained here for DEE in various representatives of Australopithecus are lower than reported previously by around 200 kcal/day. Taking into account the greater economy of human walking, particularly the effect of the longer lower limbs found in many later Homo species, also results in lowered estimates of DEE. Elongation of the lower limbs in H. erectus reduced relative travel costs nearly 50% in comparison to A.L. 288-1 (A. afarensis). The present method for calculating DEE indicates that female H. erectus DEE was 84% greater than that of female Australopithecus; this disparity is even larger than that suggested by previous workers.