A multivariate study of the Petralona skull

The Petralona skull from Greece has been generally accepted as an Upper Pleistocene variant of Neanderthal Man. It was included in a series of multivariate analyses performed by the author on cranial measurements of Pleistocene hominids. The results of the Mahalanobis D² analyses suggest that Petralona is cranially similar to the Neanderthals, but facially is distinct. Overall it resembles the Broken Hill and Djebel Ighoud skulls most closely but there are features of resemblance to Homo erectus material. The results of these analyses support recent suggestions that the Petralona material is of greater antiquity than the Upper Pleistocence.     

中国和欧洲早期智人的比较研究

中国与欧洲的早期智人头骨在颧骨额蝶突前外侧面的朝向、颧颌角、上颌骨颧突、鼻区、上面部高度、额鼻额颌缝形状、眉间区、矢状脊、印加骨和铲形门齿诸特征的形态或出现率等方面有明显差异。那时此两大地区存在相对独立的人类进化线,其间还有一定程度的基因交流。当时此两地区的人类居群分属于不同的人种。这一假说还可从古文化和古环境的资料得到支持。     

资阳人头骨化石的内部解剖结构

资阳人头骨化石,是新中国成立后首次发现的保存较为完整的人类化石,曾经引起学术界的广泛关注。由于其出土的具体位点和层位不是很清楚,对其年代目前还存有争议。长期以来,对资阳人的研究局限于传统的头骨外部形态的描述和测量。本文利用高分辨率工业CT（Computed tomography）扫描资阳人头骨,并对其内部解剖结构 (包括骨壁结构、额窦、骨内耳迷路、颞骨乳突小房、颅内模) 进行3D虚拟复原和研究,以期获得更多的形态信息。CT断层图显示资阳人头骨的骨壁板障层较厚,远大于外板层和内板层,支持以往研究认为的该头骨属于50岁以上个体的鉴定结果。额窦呈叶形,局限于两侧眼眶的内上方;左、右侧额窦的表面积分别为1780mm²和2910mm²。骨性内耳迷路半规管的大小及其比例,与尼安德特人不同,而位于晚更新世和全新世现代人变异范围之间。颞骨乳突小房为气化型,几乎占据了整个乳突部。根据复原的颅内模,资阳人的颅容量约为1250mL。颅内模的各项测量数值虽然都偏小,但脑的绝大多数形态特征、宽-高指数、顶叶-脑长指数都位于现代人的变异范围之内,而与直立人和更新世古老型人类不同。资阳人头骨内部解剖结构保留有少量原始特征,包括枕叶向后突显著、大脑窝比小脑窝大而深,这些特征不同于全新世人类,更类似于晚更新世早期现代人。     

中国古人类进化连续性新辩

本文通过对一些形态特征的分析说明研究人类起源和进化应该注意的一些思维方法的问题,并且为多地区进化说增添了新的证据:1)中国更新世头骨共同具有的眼眶及其外侧下缘形状等特征在近代头骨中已有所改变,在考虑其意义时不应该忽略时间和基因交流起过的作用;2)中国更新世头骨的共同特征并不因为它们也可见于其他地区而丧失其意义,因为证明连续性的共同特征不需要是地区独有的;3)铲形门齿的结构在近代人中变弱,不削弱它作为连续进化证据的价值,这种弱化的现象是人类进化过程中形态纤巧化的一种表现;4)在辩论时不要对有关形态作不正确的观察和曲解对方的观点和论据;5)头骨正中矢状突隆、下颌圆枕、夹紧状鼻梁和第三臼齿先天性缺失等四项特征在近代非洲人中阙如,却存在于东亚近代人,本文的分析认为这种现象增强多地区进化说的说服力。     

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.     

A new approach to mammalian cranial analysis, illustrated by examples of prosimian primates

Latex endocranial casts, which may be made without damaging the skull, reveal much of the information provided by a hemisected skull. Examination of drawings of endocasts superimposed on skulls may provide insight into the biological significance of skull and brain morphology. The high degree of cranial flexion and the globose brain shape of Daubentonia appear to be related to the functional demands of its gnawing mechanism. The broad frontal lobes of indriids are correlated with orbital orientation; differences in frontal lobe sulcal pattern suggest greater elaboration of the motor filed for the hand in indriids than in lemurids. Several features of lorisid cranial anatomy are discussed. It is suggested that, as a first approximation, increased splanchnocranial declination in small prosimians results from the necessity of accommodating relatively large eyes in a skull with a relatively small splanchnocranium.     

Archaic and modern human distal humeral morphology

The morphology of the proximal ulna has been shown to effectively differentiate archaic or premodern humans (such as Homo heidelbergensis and H. neanderthalensis) from modern humans (H. sapiens). Accordingly, the morphology of adjacent, articulating elements should be able to distinguish these two broad groups as well. Here we test the taxonomic utility of another portion of the elbow, the distal humerus, as a discriminator of archaic and modern humans. Principal components analysis was employed on a suite of log-raw and log-shape distal humeral measures to examine differences between Neandertal and modern human distal humeri. In addition, the morphological affinities of Broken Hill (Kabwe) E.898, an archaic human distal humeral fragment from the middle Pleistocene of Zambia, and five Pliocene and early Pleistocene australopith humeri were assessed. The morphometric analyses effectively differentiated the Neandertals from the other groups, while the Broken Hill humerus appears morphologically similar to modern human distal humeri. Thus, an archaic/modern human dichotomy-as previously reported for proximal ulnar morphology-is not supported with respect to distal humeral morphology. Relative to australopiths and modern humans, Neandertal humeri are characterized by large olecranon fossae and small distodorsal medial and lateral pillars. The seeming disparity in morphological affinities of proximal ulnae (in which all archaic human groups appear distinct from modern humans) and distal humeri (in which Neandertals appear distinct from modern humans, but other archaic humans do not) is probably indicative of a highly variable, possibly transitional population of which our knowledge is hampered by sample-size limitations imposed by the scarcity of middle-to-late Pleistocene premodern human fossils outside of Europe.     

Brief communication: The human humerus from the Broken Hill Mine, Kabwe, Zambia

The distal half of a right human humerus (E.898), recovered ex situ in 1925 by Hrdli?ka at the Broken Hill Mine, Kabwe, Zambia, has figured prominently in assessments of Middle Pleistocene Homo postcranial variation and of the phylogenetic polarity and functional anatomy of Pleistocene Homo upper limb morphology. Reassessment of distal humeral features that distinguish modern human and some archaic Homo humeri, especially relative olecranon breadth and medial and lateral pillar thicknesses, confirm previous studies placing it morphologically close to recent humans, as well as possibly to Early Pleistocene Homo. However, it completely lacks stratigraphic context, and there is faunal and archeological evidence for human activity at Broken Hill from the Middle Pleistocene to the Holocene. Given its uncertain geological age and modern human morphology, the Broken Hill E.898 humerus should not be used in analyses of Pleistocene humans until it is securely dated. Am J Phys Anthropol 149:312–317, 2012. © 2012 Wiley Periodicals, Inc.     

Cranial remains of Middle Pleistocene European hominids

Cranial remains of the Middle Pleistocene European hominids are reviewed in a historical context. Swanscombe and Steinheim, the earliest specimens to be discovered, are compared with each other. Following this, the more recently discovered specimens from Vértesszöllös, Petralona and Bilzingsleben are compared. Finally, Riss remains from La Chaise, Biache and Arago are discussed in the context provided by the above specimens. It is suggested that while phyletic evolution may play a role in the observed differences, a marked degree of sexual dimorphism contributes to the variation expressed by these specimens.     

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。     

Influence of the lateral ventricles and irregular skull base on brain kinematics due to sagittal plane head rotation

Two-dimensional physical models of the human head were used to investigate how the lateral ventricles and irregular skull base influence kinematics in the medial brain during sagittal angular head dynamics. Silicone gel simulated the brain and was separatedfrom the surrounding skull vessel by paraffin that provided a slip interface between the gel and vessel. A humanlike skull base model (HSB) included a surrogate skull base mimicking the irregular geometry of the human. An HSBV model added an elliptical inclusion filled with liquid paraffin simulating the lateral ventricles to the HSB model. A simplified skull base model (SSBV) included ventricle substitute but approximated the anterior and middle cranial fossae by a flat and slightly angled surface. The models were exposed to 7600 rad/s2 peak angular acceleration with 6 ms pulse duration and 5 deg forced rotation. After 90 deg free rotation, the models were decelerated during 30 ms. Rigid body displacement, shear strain and principal strains were determined from high-speed video recorded trajectories of grid markers in the surrogate brains. Peak values of inferior brain surface displacement and strains were up to 10.9X (times) and 3.3X higher in SSBV than in HSBV. Peak strain was up to 2.7X higher in HSB than in HSBV. The results indicate that the irregular skull base protects nerves and vessels passing through the cranial floor by reducing brain displacement and that the intraventricular cerebrospinal fluid relieves strain in regions inferior and superior to the ventricles. The ventricles and irregular skull base are necessary in modeling head impact and understanding brain injury mechanisms.     

Florisbad and human population succession in Southern Africa.

The human cranium recovered at Florisbad in 1932 is compared with other Sub-Saharan African hominid remains from Broken Hill, the Omo and Klasies River Mouth. The Florisbad frontal is very broad, but despite this breadth and differences in zygomatic form, there is a definite resemblance to archaic Homo sapiens from Broken Hill. There is also some similarity to both Omo I and Omo II, while fragmentary remains from Klasies River are more lightly built and hence more modern in appearance. These impressions are strengthened by measurement and statistical analysis, which demonstrates that Florisbad and Broken Hill are distant from recent African populations. Even if Florisbad is less archaic than the earlier (Middle Pleistocene?) hominid, it is not noticeably Bushman-like. New dates suggestive of early Upper Pleistocene antiquity also place Florisbad securely in a lineage containing Broken Hill, and there is no evidence to support special ties with any one group of living Africans.     

Ontogenic growth changes of the skull base in four genera of nonhuman primates.

Cross-sectional studies of the degree of the cranial base flexion were carried out in infant, juvenile and adult skulls in four genera of nonhuman primates (P. paniscus, H. lar, P. urinus, and M. mullatta). The cephalometric observations of the cranial base included linear and angular measurements of each specimen. The data obtained in this study showed that the anterior portion of the cranial base exhibits a significant shortening trend as the mammalian evolutionary scale ascends. Moreover, the growth pattern of the anterior portion of the skull base follows that of the facial bony structures. The ontogenic growth changes of the posterior portion of the skull base follows the growth pattern of the endocranial cavity. The significant trend of elongation in this area directly contributes to the posterior migration of the foramen magnum. The magnitude of these growth changes decreases as the evolutionary scale ascends. The angular measurements of the cranial flexion showed a less obtuse cranial base angle in young specimens and the ones higher on the mammalian scale. The skull kyphosis was less pronounced in these specimens and the anatomical features of the cranial base were more humanlike, including the balance of the head expressed by the position of the foramen magnum.     

福建漳平奇和洞发现的新石器时代早期人类头骨

2011年在福建漳平奇和洞发现的距今1万年左右的新石器时代早期人类遗骸"奇和洞III号",是迄今在福建地区发现的最早、最完整的古人类头骨,为探讨华南更新世晚期向全新世过渡阶段人类的体质特征及现代人群的形成与分化提供了重要的研究材料。本文对这件头骨进行了研究,奇和洞III号为35岁左右的男性个体,牙齿龋病严重,推测当时人类的经济模式主要以农耕为主。通过与更新世晚期柳江、山顶洞101号及14组新石器时代人类头骨的比较,发现奇和洞III号头骨兼有更新世晚期人类及新石器南、北方居民的混合体质特征:奇和洞III号头骨长而脑量大,似更新世晚期人类;其高而狭窄的面部、宽阔而低矮的鼻部,呈现出不同于南、北方人群的特殊体质特征。主成分分析显示,奇和洞III号与对比的新石器时代各组在头骨的测量数据上没有表现为明显的南、北地区间差异,但在头骨的测量指数或形状上存在时代和地区间的不同。本文研究为新旧石器过渡阶段人类体质特征的变异提供了进一步证据。     

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.     

内蒙古中部新发现的始施氏貘(哺乳纲,奇蹄目)头骨材料(英文)

记述了产自内蒙古呼和勃尔和剖面阿山头组的始施氏貘(Schlosseria magister)幼年头骨、头骨碎片及产自额尔登敖包底白层的S.magister成年头骨。幼年头骨在脊齿貘科属首次描述,成年头骨材料也是目前S.magister中首次描述。幼年头骨主要特征如下:头骨细长,脑颅部略有扩张,有眶后突,眶后收缩明显,矢状脊轻微发育;鼻切迹浅,位于前臼齿列之前,由前颌骨和鼻骨构成;眼眶大,眶前缘位于M1后部上方,眶下孔位于DP3-4之上;基蝶骨向后向中央逐渐加厚,末端隆起;翼蝶骨很大,从腹面看向后向背侧扩展,末端形成三角形的翼蝶骨突,覆盖在卵圆孔上;岩骨岬部表面有内颈动脉及其分支留下的3条沟痕;最内侧的为内颈动脉内侧沟,沿着岬部弯曲前行至最前部;镫骨动脉沟短小,横跨在圆窗前腹侧;岬动脉沟最长,起始于卵圆窗前内侧,沿岬部向前延伸;弓形下窝所在位置平滑,无凹陷。S.magister乳颊齿主要特征如下:DP2冠面大致呈三角形,前窄后宽,前缘较尖,长明显大于宽;外脊上仅有一个中央主尖前尖,一个非常不明显的小棱(可能为雏形的原脊)紧贴在前尖后舌侧壁上;前、后附尖不明显。DP3冠面呈梯形,与DP2相比明显增大,亚臼齿化,前附尖和后附尖略大,原、后脊明显。前尖大,后尖尚未分离;原尖很弱,几乎无法辨认,原脊低且不发育;次尖大而钝,比原尖更靠舌侧,后脊比原脊略发育,中部具小的后小尖;后脊在次尖处拐向后唇侧,使得磨蚀面呈V形。DP4冠面近方形,完全臼齿化,后尖已从外脊上分化出来,比前尖稍小,向舌侧倾斜,后尖肋明显;舌侧尖、脊发育完好,原尖和次尖大而钝,原脊、后脊近乎平行,比DP3的更高更长;两条脊分别在原尖和后尖处拐向后唇侧方,形成V形的磨蚀面。S.magister在由幼年向成年转变的过程中,主要变化趋势如下:1)吻部特征不同,主要表现为鼻切迹的位置、形态以及与之相关的前颌骨、上颌骨形态的差异。幼年头骨的鼻切迹位于前臼齿列之前,由前颌骨和鼻骨组成;成年头骨的鼻切迹后缩至M1-2之上,由鼻骨和上颌骨组成,并且因鼻切迹后缩造成鼻骨不与前颌骨接触。幼年和成年个体上颌骨的整体形态,眶前窝、眶下孔的位置和形态都差异显著。2)与咀嚼功能相关的结构改变。幼年个体的矢状脊微弱,而成年个体的则高且突起,暗示了后者具有相对强大的颞肌,以适应咀嚼功能。对比发现,S.magister与Lophialetes expeditus成年头骨在大小、整体形态和一些具有分类意义的特征上(如鼻骨和泪骨、前颌骨的接触方式,眶后突、关节后突、下颌关节窝的形状,矢状脊的高度等)非常接近。参照童永生、雷奕振(1984)对脊齿貘类头骨的划分方法,将Schlosseria magister的头骨与L.expeditus的划为一组,同时纠正了原有划分方案中存在的问题。     

大荔颅骨的测量研究

本文报道大荔颅骨的一系列测量数据, 并且将其与中国, 欧洲和非洲的中更新世人类的相应数据进行比较, 发现大荔颅骨的测量数据大多没有超出中国和欧洲/非洲中更新世人的变异范围, 有的与中国中更新世人接近, 有的与欧洲和/或非洲标本更加接近。本文将这些结果与大荔颅骨的与中国古人类共同具有的其他测量和观察特征进行综合考虑, 建议大荔人群属于中国古人类连续进化链中的一员, 并且表现出中国古人类与欧洲和非洲古人类之间基因交流的形态证据。     

中国古人类颅容量的推算方法比较

古人类的体质特征和现代人不同,依据现代人头骨测量数据计算出来的公式往往不适于古人类颅容量的推算。获取古人类颅容量最准确的方法是复原出其内部的颅内模;然而,由于颅内模的复原工艺复杂,加上古人类头骨化石数量稀少且多数残破,如何准确地推算其颅容量,成为古人学者研究的难点问题之一。本文通过对中国境内发现的不同演化阶段的古人类颅容量推算方法的对比和验证,试图找出推算古人类颅容量的最适合的公式法。研究结果显示：1）早期现代人解剖特征同现代人基本接近,依据现代人头骨推算出来的回归方程可以用来推算其颅容量;2）直立人头骨厚重、脑颅低矮,体质特征不同于现代人,其颅容量的推算不能使用现代人公式法。依据本文中国直立人头骨测量数据推导出来的回程方程（C=-1301.944+60606L+0.718b+9.936h）适合其颅容量的推算。采用此直立人公式法,推算出蓝田直立人的颅容量为918 mL;3）古老型智人的体质特征位于直立人和现代人之间,对其颅容量的推算不能一概而论：体质特征接近直立人的,如大荔人、华龙洞6号,可采用直立人公式法;体质特征接近早期现代人的,如许昌1号,可采用现代人公式法;体质特征位于直立人和早期现代人中间位置的,如马坝人、金牛山人,其颅容量约等于采用现代人公式法和采用直立人公式法获得的颅容量的平均值。     

The ‘Unicorn’ Dinosaur That Wasn’t: A New Reconstruction of the Crest of Tsintaosaurus and the Early Evolution of the Lambeosaurine Crest and Rostrum

The lambeosaurine Tsintaosaurus spinorhinus has traditionally been reconstructed with an elevated, hollow, spike-like crest composed entirely of the nasal bones, although this has been disputed. Here, we provide a new reconstruction of the skull of this species based on reexamination and reinterpretation of the morphology and articular relationships of the type and Paratype skulls and a fragmentary crest. We confirm the presence of a supracranial crest composed of the elevated nasal bones, but also including the premaxillae. We hypothesize that the crest is a tall, lobate, hollow structure that projects dorsally and slightly caudally a distance greater than the height of the skull along the quadrate. In our reconstruction, the nasal passage passes through the crest, but enters the skull rostral to the tubular process of the nasals, not through it. Tsintaosaurus spinorhinus is rediagnosed on the basis of a suite of cranial autapomorphies including a circumnarial fossa subdivided into three accessory fossae, prefrontal with ascending rostral process and lateral flange, nasals fused sagittally to form elongate tubular process that rises dorsally from skull roof, each nasal being expanded rostrocaudally into a rhomboid distal process, and medial processes of premaxillae at the summit of the cranial crest inserted between rhomboid processes of nasals. Tsintaosaurus spinorhinus lacks characters that are present in more derived lambeosaurines (parasaurolophins and lambeosaurins), such as rotation of the caudal margin of the crest to an acute angle with the skull roof, lateral processes of the nasals that enclose part of the intracranial cavity and participate in the formation of the walls of the common median chamber, and a smooth narial fossa lacking ridges and accessory fossae. We hypothesize that ancestrally the rostrum of lambeosaurines may have been more similar to that in Saurolophinae, and became subsequently reduced in complexity during evolution of the group.