Comparative myology of the hominoid cranial base. II. The muscles of the prevertebral and upper pharyngeal region

Detailed accounts of the muscular relations and attachments in the prevertebral and upper pharyngeal region of the cranial base of extant hominoids and fossil hominids are described in this paper. Marked differences exist in the cranial attachments of the longus capitis, superior constrictor, tensor palati and levator palati muscles between the three great apes and Homo sapiens. Many of the bony ridges and processes characteristic of the three great apes are also present in the 'gracile' australopithecines. Some of the differences between the muscle markings in this region of the cranial base in certain fossil hominids are discussed in the light of the findings of the dissection study of the three great apes and previous studies of the hominoid cranial base.     

Hominid cranial remains from upper Pleistocene deposits at Aduma, Middle Awash, Ethiopia

The Upper Pleistocene localities of Aduma and Bouri have yielded hominid fossils and extensive Middle Stone Age (MSA) archaeological assemblages. The vertebrate fossils recovered include parts of four hominid crania from Aduma and a complete right parietal from Bouri. Archaeological associations and radiometric techniques suggest an Upper Pleistocene age for these hominids. The more complete cranium from Aduma (ADU-VP-1/3) comprises most of the parietals, the occipital, and part of the frontal. This cranium is compared to late Middle and Upper Pleistocene hominid crania from Africa and the Middle East. The Aduma cranium shows a mosaic of cranial features shared with "premodern" and anatomically modern Homo sapiens. However, the posterior and lateral cranial dimensions, and most of its anatomy, are centered among modern humans and resemble specimens from Omo, Skhul, and Qafzeh. As a result, the Aduma and Bouri Upper Pleistocene hominids are assigned to anatomically modern Homo sapiens.     

现代中国人颞骨乳突后部的形态变异

人类颞骨因其复杂的表面及内部结构成为演化研究的重要解剖部位之一,然而由于缺乏对现代人颞骨形态与变异的细致研究及对比数据,对颞骨一些特征的定义和鉴定价值还存在争议。迄今为止,尚无学者对现代中国人群颞骨形态与变异做过专门研究。有鉴于此,本文对颞骨乳突后部一些典型性状的形态变异在现代中国人及部分古人类的表现情况进行了研究。研究结果表明:1)除乳突旁隆起的发育水平存在性别差异外(不受地区差异影响),乳突切迹、枕乳嵴、枕动脉沟的出现率和发育水平既不受地区差异影响,又无性别差异。2)现代中国人乳突后部形态总体表现为窄而深的乳突切迹、明显的乳突旁隆起、以及发育程度较弱的枕动脉沟和枕乳嵴;3)在本文所研究的性状中,乳突切迹、乳突切迹前端隆起、乳突旁隆起、枕乳嵴和枕动脉沟均呈现不同程度的个体变异;4)一些被认为属于尼安德特人衍生特征的性状在中国古人类和现代中国人的乳突后部都有出现;5)本文研究的颞骨乳突后部形态特征在中国更新世晚期人类的表现与现代中国人接近。     

Rudabánya: A late miocene subtropical swamp deposit with evidence of the origin of the African apes and humans

Rudabánya, a rich late Miocene fossil site in northern central Hungary, has yielded an abundant record of fossil primates, including the primitive catarrhine Anapithecus and the early great ape Dryopithecus. While the affinities of Anapithecus are not clear, Dryopithecus is clearly a great ape sharing numerous characteristics of its dental, cranial and postcranial anatomy with living great apes. Like all Miocene hominids (great apes and humans), Dryopithecus is more primitive in a number of ways than any living hominid, which is probably related to the passage of time since the divergence of the various lineages of living hominids, allowing for similar refinements in morphology and adaptation to take place independently. On the other hand, Dryopithecus (and Ouranopithecus) share derived characters with hominines (African apes and humans), and Sivapithecus (and Ankarapithecus) share derived characters with orangutans, thus dating the split between pongines and hominines to a time before the evolution of these fossil great apes. Pongines and hominines follow similar fates in the late Miocene, the pongines moving south into Southeast Asia from southern or eastern Asia and the hominines moving south into East Africa from the Mediterranean region, between 6 to 9 Ma.     

Estimating hominid life history: the critical interbirth interval

Unlike any great apes, humans have expanded into a wide variety of habitats during the course of evolution, beginning with the transition by australopithecines from forest to savanna habitation. Novel environments are likely to have imposed hominids a demographic challenge due to such factors as higher predation risk and scarcer food resources. In fact, recent studies have found a paucity of older relative to younger adults in hominid fossil remains, indicating considerably high adult mortality in australopithecines, early Homo, and Neanderthals. It is not clear to date why only human ancestors among all hominoid species could survive in these harsh environments. In this paper, we explore the possibility that hominids had shorter interbirth intervals to enhance fertility than the extant apes. To infer interbirth intervals in fossil hominids, we introduce the notion of the critical interbirth interval, or the threshold length of birth spacing above which a population is expected to go to extinction. We develop a new method to obtain the critical interbirth intervals of hominids based on the observed ratios of older adults to all adults in fossil samples. Our analysis suggests that the critical interbirth intervals of australopithecines, early Homo, and Neanderthals are significantly shorter than the observed interbirth intervals of extant great apes. We also discuss possible factors that may have caused the evolutionary divergence of hominid life history traits from those of great apes.     

Endocranial hyperostosis in Sangiran 2, Gibraltar 1, and Shanidar 5

Sangiran hominid 2 (S-2), Gibraltar hominid 1 (G-1), and Shanidar hominid 5 (SH-5) exhibit previously undescribed bilateral, paramedian hyperostosis of the endocranial frontal squama that spares the frontal crest, sagittal sinus, and ectocranial surface. The hyperostosis is localized to the frontal (usually the middle third) and parietal and is consistent with a diagnosis of hyperostosis calvariae interna (HCI), inclusive of hyperostosis frontalis interna. The hyperostosis in these specimens is compared to fossil hominids from Indonesia and Europe and to modern human cases of HCI. The three cases of HCI reported here document the existence and frequency of HCI in fossil hominids and push the antiquity of the disease back to nearly 1.5 millin years. The relatively great incidence of HCI in fossil hominids adds another confounding factor to the problematical issue of the taxonomic significance of cranial vault thickness. Am J Phys Anthropol 102:111–122. © 1997 Wiley-Liss, Inc.     

Foramen magnum position variation in Pan troglodytes, Plio-Pleistocene hominids, and recent Homo sapiens: implications for recognizing the earliest hominids

The anteroposterior position of the foramen magnum distinguishes living Homo sapiens from apes, and has been used as evidence for the hominid status of numerable fossils in the history of human paleontology. During the past decade, foramen magnum position has been cited as evidence of the hominid status of Ardipithecus and Sahelanthropus. Specifically, the basion of Ardpithecus is reported to be inline with the bicarotid chord, while the basion of Sahelanthropus is reported to both touch the biporion chord and intersect the bicarotid chord. In order to assess the effectiveness of anteroposterior foramen magnum position in distinguishing hominids from nonhominid apes, this study examined whether or not the positions of biporion and bicarotid relative to basion sufficiently distinguished Pan troglodytes from recent Homo sapiens and Plio-Pleistocene hominids. The distances from basion to the biporion chord (BSBIP) and from basion to the bicarotid chord (BSBIC) were measured on samples of chimpanzee (n = 69) and recent human (n = 42) crania and a sample of Plio-Pleistocene hominid fossils (n = 8). The data were used to test the hypothesis that BSBIP and BSBIC measurements do not sufficiently distinguish P. troglodytes from hominids. While basion to biporion (BSBIP) does not effectively distinguish P. troglodytes from Plio-Pleistocene hominids and humans when used univariately, basion to bicarotid (BSBIC), when used univariately or bivariately with BSBIP, can be used to test whether or not an unknown specimen is a hominid. These results are used to evaluate the hominid status of Ardipithecus and Sahelanthropus.     

Barking up the wrong ape--australopiths and the quest for chimpanzee characters in hominid fossils

With the shift during the 1980s from a human-great ape ultimately to an orangutan-(gorilla-(human-chimp)) theory of relatedness, the search for chimpanzee-like features in early hominids intensified. Reconstructions of early hominids became caricatures of chimpanzees, not only in soft tissue features (e.g. the nasal region), but in supposed bony structures (e.g. an anteriorly and especially superiorly protruding a supraorbital torus with a distinct posttoral sulcus behind). In spite of rampant >Panophilia,< actual morphologies of the majority of early hominid specimens are those cited as uniting an orangutan clade. Those specimens that are >chimpanzee-like< are probably not cladistically hominid.     

化石人类脑演化研究概况

脑演化是人类演化的一个重要组成部分,其研究可以为人类起源、演化、人群关系及语言、智力等方面提供重要的信息。脑演化的主要证据是通过研究颅内模（endocast）及颅骨的形态得到的。颅内模是从颅骨内表面得到的脑的外部形态。有时颅骨的内腔充满泥沙,并且为钙质所结固,可以自然形成颅内模。也可以人工制作颅内模。颅内模和颅骨的内表面能够提供人类脑髓及神经进化方面的直接证据。对化石人类脑演化的研究主要包括以下几个方面的内容：测量或估计脑量的大小及其和身体大小之间的关系;研究脑量随时间的变化过程;通过对早期人类颅内模表面沟回形态特征的研究,探索脑功能区在早期人类和猿类的区别及在演化上的变化;左右大脑不对称性与一侧优势关系,探讨语言的起源和惯用手的脑功能基础等;脑膜中动脉系统、静脉窦系统及与血液循环相关的排泄孔的变化,探讨大脑各部分比例的变化和功能的日益复杂对供血需求的影响;通过对人类脑演化的研究,探讨人类进化的原因。本文通过对以上几个方面及其中国化石脑演化研究的介绍,对化石人类脑演化的研究概况作综合论述和简单回顾。     

Cranial base morphology and temporal bone pneumatization in Asian Homo erectus

The external morphological features of the temporal bone are used frequently to determine taxonomic affinities of fossils of the genus Homo. Temporal bone pneumatization has been widely studied in great apes and in early hominids. However, this feature is rarely examined in the later hominids, particularly in Asian Homo erectus. We provide a comparative morphological and quantitative analysis of Asian Homo erectus from the sites of Ngandong, Sambungmacan, and Zhoukoudian, and of Neandertals and anatomically modern Homo sapiens in order to discuss causes and modalities of temporal bone pneumatization during hominid evolution. The evolution of temporal bone pneumatization in the genus Homo is more complex than previously described. Indeed, the Zhoukoudian fossils have a unique pattern of temporal bone pneumatization, whereas Ngandong and Sambungmacan fossils, as well as the Neandertals, more closely resemble the modern human pattern. Moreover, these Chinese fossils are characterized by a wide midvault and a relatively narrow occipital bone. Our results support the point of view that cell development does not play an active role in determining cranial base morphology. Instead, pneumatization is related to available space and to temporal bone morphology, and its development is related to correlated morphology and the relative disposition of the bones and cerebral lobes. Because variation in pneumatization is extensive within the same species, the phyletic implications of pneumatization are limited in the taxa considered here.     

An alternative method of estimating the cranial capacity of Olduvai Hominid 7

The cranial capacity of Olduvai Hominid 7 is estimated to be 690 cc, with a standard uncertainty range of 538 to 868 cc. The estimate is derived from a systematic consideration of the relationships between Bregma-Asterion chords and cranial capacities obtained from a large sample of Homo sapiens and Pan troglodytes and from available fossil hominids. The estimation technique is applicable to other characters and specimens.     

The ATD6-5 mandibular specimen from Gran Dolina (Atapuerca, Spain). Morphological study and phylogenetic implications.

Metric and shape features of the Lower Pleistocene mandibular specimen ATD605 from the level 6 of Gran Dolina site (Atapuerca, Spain) are compared with a large sample of fossil hominid mandibles. The analysis shows that ATD6-5 displays a generalized morphology largely shared with both African and European Lower and Middle Pleistocene samples. However, distinctive African traits, such as corpus robustness and strong alveolar prominence, are absent in the Gran Dolina specimen. At the same time, none of the apomorphic features that characterize Middle and early Upper Pleistocene European hominids can be recognized in ATD6-5. Finally, the Gran Dolina specimen displays a remarkable position of the mylohyoid groove, only comparable to that found in immature specimens of Homo ergaster, and very rarely in adult H. sapiens. The morphology of ATD6-5 supports the hypothesis of an African origin for the first Europeans with subsequent phylogenetic continuity with Middle Pleistocene populations in Europe. These findings are consistent with H. antecessor being the last common ancestor of Neandertals and H. sapiens.     

The posterior border of the sphenoid greater wing and its phylogenetic usefulness in human evolution

The elucidation of patterns of cranial skeletal maturation and growth in fossil hominids is possible not only through dental studies but also by mapping different aspects of ossification in both extant African apes and humans. However, knowledge of normal skeletal development in large samples of extant great apes is flimsy. To remedy this situation, this paper offers an extensive survey and thorough discussion of the ossification of the posterior border of the sphenoid greater wing. Indeed, this area provides much information about basicranial skeletal maturation. We investigate three variants: the absence of the foramen spinosum and the position of both the foramen spinosum and the foramen ovale in relation to the sphenosquamosal suture. Providing original data about humans and 1,425 extant great ape skulls and using a sample of 64 fossil hominids, this study aimed to test whether different ossification patterns occurred during the course of human evolution. The incidence of three derived morphologies located on the posterior border of the sphenoid greater wing increases during human evolution at different geological periods. The evolutionary polarity of these three derived morphologies is assessed by outgroup comparison and ontogenetic methods. During human evolution, there is a clear trend for the foramen spinosum to be present and wholly located on the posterior area of the sphenoid greater wing. Moreover, in all the great ape species and in Australopithecus afarensis, the sphenosquamosal suture may split the foramen ovale. Inversely, the foramen ovale always lies wholly within the sphenoid greater wing in Australopithecus africanus, robust australopithecines, early Homo, H. erectus (and/or H. ergaster), and Homo sapiens. From ontogenetic studies in humans, we conclude that, during human evolution, the ossification of the posterior area of the sphenoid greater wing progressively surrounded the middle meningeal artery (passing through the foramen spinosum) and the small meningeal artery (passing through the foramen ovale). Am J Phys Anthropol 107:387–399, 1998. © 1998 Wiley-Liss, Inc.     

The eruption pattern of the permanent incisors and first permanent molars in Australopithecus (Paranthropus) robustus

This study aims to reassess the claim that the eruption sequence of the permanent incisor and first permanent molar teeth of Australopithecus (Paranthropus) robustus is identical with that in modern Homo sapiens. Eight fossil hominid mandibles of equivalent dental developmental age were chosen for comparative study. Emphasis has been placed upon the comparative timing of events within the growth period rather than eruption sequence alone. The results of this study indicate that Homo sapiens and Australopithecus (Paranthropus) robustus share the same pattern of permanent molar and incisor eruption and that this is significantly different from the pattern of eruption shared by the great apes, Australopithecus africanus and Australopithecus afarensis.     

Meningeal arterial patterns in great apes: Implications for hominid vascular evolution

Arterial meningeal patterns were observed for 100 hemispheres from great ape endocasts (Pan paniscus, Pan troglodytes, Gorilla gorilla, and Pongo pygmaeus). Eight patterns emerged based on the relative contributions to the walls and dura mater of the middle part of the braincase of meningeal arteries that stem from two sources. These arteries enter the braincase through either the orbit (delivering blood from the internal carotid artery) or through the base of the middle cranial fossa (via the middle meningeal artery whose blood comes from the external carotid artery). The three genera of apes manifest different frequencies of the eight, patterns, with orangutans highly dependent on orbital meningeal arteries at one extreme, and chimpanzees showing the greatest reliance on the middle meningeal artery at the other. As was the case in an earlier study of rhesus monkeys, there is a trend across the two genera of African apes for increased mean cranial capacity to be associated with increased reliance on the internal carotid artery for supplying the middle portion of the braincase. However, unlike the case for macaques, this trend does not reach statistical significance in African apes. Because it is rare for humans to manifest significant arterial contributions from the orbit to the middle cranial fossa, the comparative data on monkeys, apes, and humans suggest that, during the course of vascular evolution in Homo, the middle meningeal artery eventually took over supply of the entire middle cranial fossa. This hypothesis should be tested in the hominid fossil record. Earlier work on meningeal arterial patterns in apes has traditionally relied on Adachi's system that was determined from humans and focuses on the origin of the middle branch of the middle meningeal artery. As a result, the extensive orbital contributions to the middle portion of the braincase that characterize apes were not recognized and the eight patterns described in this paper were often erroneously assigned to the three patterns that adequately describe only humans. Adachi's system should therefore be abandoned for nonhuman primates and early hominids. A correct understanding of meningeal arterial evolution cannot be achieved until the orbital contributions to the meningeal arteries are recognized and incorporated into an evolutionary study that spans from apes to fossil hominids to living people. © 1993 Wiley-Liss, Inc.     

Surgical anatomy of the internal carotid artery

Starting from the carotid trigone, a surgical approach to the parapharyngeal part of the internal carotid artery is described. The retrostyloidal part of the lateropharyngeal space is opened up from laterocaudal after resecting the posterior belly of the digastric muscle and the styloidal muscles. Vulneration of the cranial nerves (VII, IX, X) has to be prevented.     

Developmental age and taxonomic affinity of the Mojokerto child,Java, Indonesia

An increasing number of claims place hominids outside Africa and deep in Southeast Asia at about the same time that Homo erectus first appears in Africa. The most complete of the early specimens is the partial child's calvaria from Mojokerto (Perning I), Java, Indonesia. Discovered in 1936, the child has been assigned to Australopithecus and multiple species of Homo, including H. modjokertensis, and given developmental ages ranging from 1–8 years. This study systematically assesses Mojokerto relative to modern human and fossil hominid growth series and relative to adult fossil hominids. Cranial base and vault comparisons between Mojokerto and H. sapiens sapiens (Hss) (n = 56), Neandertal (n = 4), and H. erectus (n = 4) juveniles suggest a developmental age range between 4 and 6 years. This range is based in part on new standards for assessing the relative development of the glenoid fossa. Regression analyses of vault arcs and chords indicate that H. erectus juveniles have more rounded frontals and less angulated occipitals than their adult counterparts, whereas Hss juveniles do not show these differences relative to adults. The growth of the cranial superstructures and face appear critical to creating differences in vault contours between H. erectus and Hss. In comparison with adult H. erectus and early Homo (n = 27) and adult Hss (n = 179), the Mojokerto child is best considered a juvenile H. erectus on the basis of synapomorphies of the cranial vault, particularly a metopic eminence and occipital torus, as well as a suite of characters that describe but do not define H. erectus, including obelion depression, supratoral gutter, postorbital constriction, mastoid fissure, lack of sphenoid contribution to glenoid fossa, and length and breadth ratios of the temporomandibular joint. Mojokerto is similar to other juvenile H. erectus in the degree of development of its cranial superstructures and its vault contours relative to adult Indonesian specimens. The synapomorphies which Mojokerto shares with H. erectus are often considered autapomorphies of Asian H. erectus and confirm the early establishment and long-term continuity of the Asian H. erectus bauplan. This continuity does not, however, necessarily reflect on the pattern of origin of modern humans in the region. Am J Phys Anthropol 102:497–514, 1997. © 1997 Wiley-Liss, Inc.