Asymmetric vault modification in Hopi crania

Cradleboarding was practiced by numerous prehistoric and historic populations, including the Hopi. In this group, one result of cra-dleboarding was bilateral or asymmetric flattening of the posterior occipital. We test whether cradleboarding had significant effects on the morphology of the cranial vault, cranial base, and face. Additionally, we examine associations between direction of flattening and asymmetric craniofacial growth. A skeletal sample of Hopi from the Old Walpi site includes both nonmodified (N = 43) and modified individuals (N = 39). Three-dimensional coordinates of 53 landmarks were obtained using a diagraph. Thirty-six landmarks were used to define nine finite elements in the cranial vault, cranial base, and face. Finite element scaling was used to compare average nonmodified individuals, with averages of bilaterally, right, and left modified individuals. The significance of variation among “treatment” groups was evaluated using a bootstrap test. Pearson product-moment correlations test the association of asymmetry with direction of modification. Hopi cradleboarding has a significant effect on growth of the cranial vault, but does not affect morphology of the cranial base or face. Bilateral flattening of the cranial vault leads to decreased length and increased width of the cranial vault. Flattening of the right or left cranial vault results in ipsilaterally decreased length and width coupled with a corresponding increased length and width on the contralateral side of the cranial vault. There is a significant correlation of size asymmetry with direction of modification in the cranial vault, but not with size or shape change in the cranial base or face. © 1995 Wiley-Liss, Inc.     

The Developmental Basis of Quantitative Craniofacial Variation in Humans and Mice

The human skull is a complex and highly integrated structure that has long held the fascination of anthropologists and evolutionary biologists. Recent studies of the genetics of craniofacial variation reveal a very complex and multifactorial picture. These findings contrast with older ideas that posit much simpler developmental bases for variation in cranial morphology such as the growth of the brain or the growth of the chondrocranium relative to the dermatocranium. Such processes have been shown to have major effects on cranial morphology in mice. It is not known, however, whether they are relevant to explaining normal phenotypic variation in humans. To answer this question, we obtained vectors of shape change from mutant mouse models in which the developmental basis for the craniofacial phenotype is known to varying degrees, and compared these to a homologous dataset constructed from human crania obtained from a single population with a known genealogy. Our results show that the shape vectors associated with perturbations to chondrocranial growth, brain growth, and body size in mice do largely correspond to axes of covariation in humans. This finding supports the view that the developmental basis for craniofacial variation funnels down to a relatively small number of key developmental processes that are similar across mice and humans. Understanding these processes and how they influence craniofacial shape provides fundamental insights into the developmental basis for evolutionary change in the human skull as well as the developmental-genetic basis for normal phenotypic variation in craniofacial form.     

Chondrocranial development in larval Rana sylvatica (Anura: Ranidae): morphometric analysis of cranial allometry and ontogenetic shape change

This study provides baseline quantitative data on the morphological development of the chondrocranium in a larval anuran. Both linear and geometric morphometric methods are used to quantitatively analyze size-related shape change in a complete developmental series of larvae of the wood frog, Rana sylvatica. The null hypothesis of isometry was rejected in all geometric morphometric and most linear morphometric analyses. Reduced major axis regressions of 11 linear chondrocranial measurements on size indicate a mixture of allometric and isometric scaling. Measurements in the otic and oral regions tend to scale with negative allometry and those associated with the palatoquadrate and muscular process scale with isometry or positive allometry. Geometric morphometric analyses, based on a set of 11 chondrocranial landmarks, include linear regression of relative warp scores and multivariate regression of partial warp scores and uniform components on log centroid size. Body size explains about one-quarter to one-third of the total shape variation found in the sample. Areas of regional shape transformation (e.g., palatoquadrate, otic region, trabecular horns) are identified by thin-plate spline deformation grids and are concordant with linear morphometric results. Thus, the anuran chondrocranium is not a static structure during premetamorphic stages and allometric patterns generally follow scaling predictions for tetrapod cranial development. Potential implications regarding larval functional morphology, cranial development, and chondrocranial evolution in anurans are discussed.     

Effects of fronto-occipital artificial cranial vault modification on the cranial base and face.

Artificial reshaping of the cranial vault has been practiced by many human groups and provides a natural experiment in which the relationships of neurocranial, cranial base, and facial growth can be investigated. We test the hypothesis that fronto-occipital artificial reshaping of the neurocranial vault results in specific changes in the cranial base and face. Fronto-occipital reshaping results from the application of pads or a cradle board which constrains cranial vault growth, limiting growth between the frontal and occipital and allowing compensatory growth of the parietals in a mediolateral direction. Two skeletal series including both normal and artificially modified crania are analyzed, a prehistoric Peruvian Ancon sample (47 normal, 64 modified crania) and a Songish Indian sample from British Columbia (6 normal, 4 modified). Three-dimensional coordinates of 53 landmarks were measured with a diagraph and used to form 9 finite elements as a prelude to finite element scaling analysis. Finite element scaling was used to compare average normal and modified crania and the results were evaluated for statistical significance using a bootstrap test. Fronto-occipitally reshaped Ancon crania are significantly different from normal in the vault, cranial base, and face. The vault is compressed along an anterior-superior to posterior-inferior axis and expanded along a mediolateral axis in modified individuals. The cranial base is wider and shallower in the modified crania and the face is foreshortened and wider with the anterior orbital rim moving inferior and posterior towards the cranial base. The Songish crania display a different modification of the vault and face, indicating that important differences may exist in the morphological effects of fronto-occipital reshaping from one group to another.     

Effects of annular cranial vault modification on the cranial base and face

Artificial modification of the cranial vault was practiced by a number of prehistoric and protohistoric populations, frequently during an infant's first year of life. We test the hypothesis that, in addition to its direct effects on the cranial vault, annular cranial vault modification has a significant indirect effect on cranial base and facial morphology. Two skeletal series from the Pacific Northwest Coast, which include both nonmodified and modified crania, were used: the Kwakiutl (62 nonmodified, 45 modified) and Nootka (28 nonmodified, 20 modified). Three-dimensional coordinates of 53 landmarks were obtained using a diagraph, and 36 landmarks were used to define nine finite elements in the cranial vault, cranial base, and face. Finite element scaling was used to compare average nonmodified and average modified crania, and the significance of the results were evaluated using a bootstrap test. Annular modification of the cranial vault produces significant effects on the morphology of the cranial base and face. Annular modification in the Kwakiutl resulted in restrictions of the cranial vault in the medial-lateral and superior-inferior dimensions and an increase in anterior-posterior growth. Similar dimensional changes are observed in the cranial base. The Kwakiutl face is increased anterior-posteriorly and reduced anterior-laterally to posterior-medially. Similar effects of modification are observed in the Nootka cranial vault and cranial base, though not in the face. These results demonstrate the developmental interdependence of the cranial vault, cranial base, and face. © 1993 Wiley-Liss, Inc.     

Development and tissue origins of the mammalian cranial base

The vertebrate cranial base is a complex structure composed of bone, cartilage and other connective tissues underlying the brain; it is intimately connected with development of the face and cranial vault. Despite its central importance in craniofacial development, morphogenesis and tissue origins of the cranial base have not been studied in detail in the mouse, an important model organism. We describe here the location and time of appearance of the cartilages of the chondrocranium. We also examine the tissue origins of the mouse cranial base using a neural crest cell lineage cell marker, Wnt1-Cre/R26R, and a mesoderm lineage cell marker, Mesp1-Cre/R26R. The chondrocranium develops between E11 and E16 in the mouse, beginning with development of the caudal (occipital) chondrocranium, followed by chondrogenesis rostrally to form the nasal capsule, and finally fusion of these two parts via the midline central stem and the lateral struts of the vault cartilages. X-Gal staining of transgenic mice from E8.0 to 10 days post-natal showed that neural crest cells contribute to all of the cartilages that form the ethmoid, presphenoid, and basisphenoid bones with the exception of the hypochiasmatic cartilages. The basioccipital bone and non-squamous parts of the temporal bones are mesoderm derived. Therefore the prechordal head is mostly composed of neural crest-derived tissues, as predicted by the New Head Hypothesis. However, the anterior location of the mesoderm-derived hypochiasmatic cartilages, which are closely linked with the extra-ocular muscles, suggests that some tissues associated with the visual apparatus may have evolved independently of the rest of the “New Head”.     

Craniofacial suture stenosis: morphologic effects

Craniofacial anomalies, such as Apert's and Crouzon's syndromes, are presumed to be related to premature growth arrest of cranial base growth sites. However, premature growth arrest at cranial vault sutures in animals appears to play a causative role in the development of cranial deformities characteristic of single-suture, or simple, craniosynostosis in humans. To study the possible causative role of cranial vault and other (interface) suture stenoses on the development of craniofacial deformity, a vault suture and an interface suture between the cranial vault and facial skeleton were simultaneously immobilized. Thirty-one New Zealand White rabbits at 9 days of age underwent implantation of dental amalgam growth markers adjacent to cranial vault and facial sutures. In the experimental group (n = 15), methylcyanoacrylate adhesive was applied over the coronal (vault) and frontonasal (interface suture between vault and facial skeleton) sutures to immobilize them. The remaining 16 animals served as sham-treated controls. All animals underwent serial radiographic cephalometry to document growth effects in the cranial vault, cranial base, and facial skeleton. Application of adhesive resulted in statistically significant (p less than 0.05) reduction in growth at the coronal and frontonasal sutures. This was accompanied by an overall significant reduction in neurocranial vault length during the first 30 days of development.(ABSTRACT TRUNCATED AT 250 WORDS)     

Cranial integration in Homo: singular warps analysis of the midsagittal plane in ontogeny and evolution

This study addresses some enduring issues of ontogenetic and evolutionary integration in the form of the hominid cranium. Our sample consists of 38 crania: 20 modern adult Homo sapiens, 14 sub-adult H. sapiens, and four archaic Homo. All specimens were CT-scanned except for two infant H. sapiens, who were imaged by MR instead. For each specimen 84 landmarks and semi-landmarks were located on the midsagittal plane and converted to Procrustes shape coordinates. Integration was quantified by the method of singular warps, a new geometric-statistical approach to visualizing correlations among regions. The two classic patterns of integration, evolutionary and ontogenetic, were jointly explored by comparing analyses of overlapping subsamples that span ranges of different hypothetical factors. Evolutionary integration is expressed in the subsample of 24 adult Homo, and ontogenetic integration in the subsample of 34 H. sapiens. In this data set, vault, cranial base, and face show striking and localized patterns of covariation over ontogeny, similar but not identical to the patterns seen over evolution. The principal differences between ontogeny and phylogeny pertain to the cranial base. There is also a component of cranial length to height ratio not reducible to either process. Our methodology allows a separation of these independent processes (and their impact on cranial shape) that conventional methods have not found.     

The transient expression of type II collagen at tissue interfaces during mammalian craniofacial development

Using immunocytochemical techniques, the spatiotemporal distribution of the major collagen isoform of cartilage, type II collagen, has been investigated during early craniofacial development in the mouse embryo. Early and transient expression was associated with the otic and optic vesicles, the ventrolateral surfaces of the developing brain, olfactory conchi, endocardial and mesocardial tissues, the lateral and basal surfaces of the pharyngeal endoderm and beneath the ectoderm of the branchial arches. A number of these locations are sites of epithelial-mesenchymal tissue interaction believed to generate the component parts of the chondrocranium; here, type II collagen appears transiently in advance of overt chondrogenesis in the mesenchyme. At such sites, immunofluorescence is typically localised along the basal surface of the epithelial partner, with the strongest reaction detected between the basal aspects of the otic and rhombencephalic epithelia. Immunoelectron microscopy, using pre-embedding immunostaining and a protein G-gold technique, reveals that the type II collagen is adjacent to, but not integral with, the basal laminae. Gold particles are clearly associated with 10-15 nm fibrils of the extracellular matrix in the reticulate lamina region. The pattern of type II collagen expression in the mouse closely correlates with that demonstrated previously in the quail, indicating a high degree of phylogenetic conservation between these two vertebrate species. These findings are consistent with the hypothesis that the pattern of epithelial secretion of type II collagen, or a coexpressed matrix molecule, constitutes a morphogenetic signal, realised as a matrix-mediated tissue interaction, and specifying the form of the vertebrate chondrocranium. Three-dimensional reconstruction of early type II collagen distribution, and of the subsequent chondrocranial cartilages, reveals that chondrocranial form can be derived from a 'pre-pattern' of epithelially derived type II collagen expressed at epithelial-mesenchymal tissue interfaces.     

Basicranial influence on overall cranial shape

This study examines the extent to which the major dimensions of the cranial base (maximum length, maximum breadth, and flexion) interact with brain volume to influence major proportions of the neurocranium and face. A model is presented for developmental interactions that occur during ontogeny between the brain and the cranial base and neurocranium, and between the neurobasicranial complex (NBC) and the face. The model is tested using exocranial and radiographic measurements of adult crania sampled from five geographically and craniometrically diverse populations. The results indicate that while variations in the breadth, length and flexion of the cranial base are mutually independent, only the maximum breadth of the cranial base (POB) has significant effects on overall cranial proportions, largely through its interactions with brain volume which influence NBC breadth. These interactions also have a slight influence on facial shape because NBC width constrains facial width, and because narrow-faced individuals tend to have antero-posteriorly longer faces relative to facial breadth than wide-faced individuals. Finally, the model highlights how integration between the cranial base and the brain may help to account for the developmental basis of some morphological variations such as occipital bunning. Among modern humans, the degree of posterior projection of the occipital bone appears to be a consequence of having a large brain on a relatively narrow cranial base. Occipital buns in Neanderthals, who have wide cranial bases relative to endocranial volume, may not be entirely homologous with the morphology occasionally evident in Homo sapiens.     

Testing hypotheses about tinkering in the fossil record: the case of the human skull

Efforts to test hypotheses about small-scale shifts in development (tinkering) that can only be observed in the fossil record pose many challenges. Here we use the origin of modern human craniofacial form to explore a series of analytical steps with which to propose and test evolutionary developmental hypotheses about the basic modules of evolutionary change. Using factor and geometric morphometric analyses of craniofacial variation in modern humans, fossil hominids, and chimpanzee crania, we identify several key shifts in integration (defined as patterns of covariation that result from interactions between components of a system) among units of the cranium that underlie the unique shape of the modern human cranium. The results indicate that facial retraction in modern humans is largely a product of three derived changes: a relatively longer anterior cranial base, a more flexed cranial base angle, and a relatively shorter upper face. By applying the Atchley-Hall model of morphogenesis, we show that these shifts are most likely the result of changes in epigenetic interactions between the cranial base and both the brain and the face. Changes in the size of the skeletal precursors to these regions may also have played some role. This kind of phenotype-to-genotype approach is a useful and important complement to more standard genotype-to-phenotype approaches, and may help to identify candidate genes involved in the origin of modern human craniofacial form.     

A factor analysis of cranial base and vault dimensions in children

Lengths within the cranial base and vault were measured in cephalometric radiographs of 220 boys and 177 girls ranging in age from 0 to 15 years; all these children are participants in The Fels Longitudinal Growth Study. The present study is based on mixed longitudinal data derived from 1640 radiographs for boys and 1260 radiographs for girls. Factor analysis was applied separately for boys and girls for each age group; i.e., 0–3, 4–6, 7–9, 10–12, and 13–15 years. For the 0–3 year age group, two factors were extracted in each sex, whereas four factors were extracted in the rest of the age groups. The factor structures are similar in the three older age groups of boys (7–9, 10–12, and 13–15 years). The first four factors for these groups are labelled, respectively: cranial vault size, posterior cranial base length, presphenoid length, and basisphenoid length. The order of the third and fourth factors is reversed in the 7–9 year olds. For girls, the factors extracted were also the same in both the 7–9 and 10–12 year age groups, even though the order of factors was different between age groups; i.e., anterior cranial base length, cranial vault size, basisphenoid length, and basioccipital length. Differential growth rates among cranial base dimensions probably cause changes in factor patterns. Obliteration of the spheno-occipital synchondrosis is suggested as the mechanism responsible for the change of factor pattern in the girls. Closure of this synchondrosis would have occurred too late to affect the patterns in boys.     

Effects of fronto-occipital cranial reshaping on mandibular form.

Cultural reshaping (artificial deformation or modification) of the neurocranial vault provides an artificially increased range of morphological variability within which the relationship between the growing neurocranium and face can be investigated. We analyze crania which have been fronto-occipitally compressed to ascertain possible morphological effects on the mandible. We collected measures of mandibular breadth, length, and height from 82 modified (N = 48) and unmodified (N = 34) crania from a Peruvian Ancon series. Angle classification was also scored in order to investigate whether or not occlusal relationships were affected by neurocranial reshaping. Only intercondylar distance (posterior mandibular breadth) exhibited significant differences between unmodified and modified groups, though this difference was relatively small compared with vault deformation. The modified crania had a higher frequency of normal occlusion (Class I) than the unmodified crania. Increased intercondylar breadth in modified skulls is due to a cascade of effects which begin with a direct effect of the fronto-occipital deforming device on neurocranial shape (increased neurocranial width). The increase in mandibular breadth may be a compensatory response to increased cranial base breadth and maintains articulation between the cranial base and mandible. The increased posterior breadth, coupled with a slight decrease in mandibular depth, may contribute to the change in occlusal relationships suggested for this sample.     

华北人欧洲人和澳洲土著人的头骨厚度

The thickness of the cranial vault at the midline on the mid-frontal squama, pre-bregmatic einence, frontal at bregma, parietal at vertex, occipital at lambda and the external occipita1 protuberance was recorded in 40 male and 7 female Northern Chinese crania, 47 male and 52 female Australian Aboriginal crania and 13 male European crania using specially nodified vernier calipers. Comparison of vault thickness data obtained through direct measurement with those obtained fron lateral radiographs indicated that direct measurenent provided consistently more accurate results.
Male and fermale samples were processed separately so that the extent of sexbased variation could be examined.Student's t test was used to compare the sample means and the percentage of sexual dimorphism for each dimension was calculated according to Garn et al, (1964).The possibility of an allometric association between the thickness of the bones within the cranial vault, size of the cranial vault and stature was examined using Spearman's rank correlation coefficient and the Australian Aboriginal sample.
All but one of the mean thickness dimensions in the Australian Aboriginal male sample is significantly greater than the Northern Chinese and European means. The female results support those obtained with the males.In both males and females thickness at the external occipital protuberance, in all of the populations examined,did not correlate highly with that obtained from other parts of the cranial vault.This reflects the high degree of morphological variation in the position of the internal occipital protuberance and its influence on cranial vault thickness dimensions recorded at the external occipital protuberance.The European and Northern Chinese samples have similar cranial vault thickness dimensions. The Spearman's rank correlation coefficient matrix scores provide sone support for a biological association between vault thickness and overall cranial size. However, there appears to be little support for an association between stature and cranial vault thickness. The difference between the male and female mean vault thickness dimensions were significant at bregma, vertex and the external occipital protuberance in Australian Aboriginals and lambda and the external occipital protuberance in Northern Chinese. Some caution is needed in the interpretation of the Northern Chinese female data as the sample is extremely small.
Evidence of trauma, supressed fractures, is extremely common on the vaults of Australian Aboriginal crania from southern and central Australia. Traditionally Australian Aboriginals, males and females, involved in agressive dispute will use a substantial wooden implement and strike to the head of thir opponent(Meggitt 1962).The injuries that result from this are more common in females than in male. This form of social interaction must have rigorously selected against those individuals with thinner bones in their cranial vaults. To a large degree this may explain the greatly thickened vaults in Australian Aboriginals relative to Europeans and Northern Chines.This may also provide a clue to the factors resulting in the development of marked cranial vault thickness in Homo erectus.
     

Cranial base and jaw relationship

The lateral skull radiographs of 124 boys aged approximately 10 years divided equally between the four angle classes were digitized in an effort to establish the relationship between cranial base size and shape and jaw relationship. Comparison of the means for occlusal groups showed a trend from class II to class III as cranial base dimensions and angle decreased. The condyle was also more distally positioned with respect to nasion, point A and the Pterygomaxillary vertical in the class II groups. Cranial base length correlated strongly with maxillary length but weakly with mandibular length. Nevertheless, the size of the maxilla did not influence its prognathism. The cranial base angle was strongly correlated (-0.7) with angle sella-nasion-point B. It is concluded that cranial base size and shape influence mandibular prognathism by determining the anteroposterior position of the condyle relative to the facial profile.     

Anthropoid cranial base architecture and scaling relationships

This paper examines how various measures of basicranial length and cranial base angulation affect the relationship between basicranial flexion and relative brain size in anthropoids, including Homo sapiens. Most recent studies support the "spatial packing" hypothesis, that basicranial flexion in haplorhines maximizes braincase volume relative to basicranial length. However, a few studies find the basicranium is less flexed in H. sapiens than expected for other anthropoids, suggesting that other factors contribute to variation in hominin basicranial flexion. The measure of relative brain size used to test the spatial packing hypothesis, the Index of Relative Encephalization (IRE), is calculated with basicranial length (BL) in its denominator, so that shorter BL and larger brain size potentially inflate H. sapiens IREs. To investigate this problem, the lengths of midline cranial floor sections were scaled relative to the cube root of endocranial volume in 157 specimens from 18 anthropoid species. Results indicate that the posterior cranial base and planum sphenoideum are significantly shorter in H. sapiens than in other anthropoids, accounting for higher IREs. Including the cribriform plate in BL, advisable in studies using anthropoids, affects whether H. sapiens differs from other anthropoids for basicranial flexion vs. IRE. However, despite a shorter BL and elevated IRE, H. sapiens does not deviate significantly from the anthropoid relationship between basicranial flexion and relative brain size for two cranial base angles. Because different measures of cranial base angulation change how H. sapiens falls along the anthropoid regression line, it remains equivocal whether the basicranium is less flexed in H. sapiens than in other anthropoids when compared to relative brain size.     

A morphometric analysis of craniofacial growth and changes in spatial relations during secondary palatal development in human embryos and fetuses

Staged human embryos and fetuses in the Carnegie Embryological Collection were morphometrically analyzed to show craniofacial dimensions and changes in spatial relations, and to identify patterns that would reflect normal developmental events during palatal formation. Normal embryos aged 7-8 weeks postconception (Streeter-O'Rahilly stages 19-23) and fetuses aged 9-10 weeks postconception, in eight groups with mean crown-rump (CR) lengths of 18-49 mm, were studied with cephalometric methods developed for histologic sections. In the 4-week period studied, facial dimensions increased predominantly in the sagittal plane with extensive changes in length (depth) and height, but limited changes in width. Growth of the mandible was more rapid than the nasomaxillary complex, and the length of Meckel's cartilage exceeded the length of the oronasal cavity at the time of horizontal movement of the shelves during stage 23. Simultaneously with shelf elevation, the upper craniofacial complex lifted, and the tongue and Meckel's cartilage extended forward beneath the primary palate. Analysis of spatial relations in the oronasal cavity showed that the palatomaxillary processes became separated from the tongue--mandibular complex as the head extended, and the tongue became positioned forward with growth of Meckel's cartilage. As the head position extended by 35 degrees, the cranial base angulation was unchanged and the primary palate maintained a 90 degrees position to the posterior cranial base. However, the sagittal position of the maxilla relative to the anterior cranial base increased by 20 degrees between stages 19 and 23. In the late embryonic and early fetal periods, the mean cranial base angulation of approximately 128 degrees and the mean maxillary position angulation of approximately 84 degrees were similar to the angulations previously shown to be present later prenatally and post-natally. The results suggest that human patterns of cranial base angulation and maxillary position to the cranial base develop during the late embryonic period when the chondrocranium and Meckel's cartilage form the primary skeleton.     

南京1号直立人头骨与肯尼亚KNM-ER 3733人类头骨化石的形态比较

KNM-ER 3733人类头骨化石的年代为距今1.78百万年,1975年发现于肯尼亚。Walker和Leakey注意到这具头骨与周口店直立人的在脑颅形态上很相近,但二者在年代上相差大约1百万年,故认为直立人形态在这1百万年期间是稳定的。长期来此观点缺乏更多的人类化石证据来支持。1993年在中国发现了南京1号人类头骨化石。该头骨与KNM-ER 3733头骨一样兼具脑颅和面颅,且都属于成年女性个体,但南京1号人类头骨化石的年代比KNM-ER 3733人类头骨化石的要晚大约1百万年。因此,南京1号人类头骨是目前所知的可用来验证直立人头骨形态是否在1百万年期间保持稳定的唯一合适的人类头骨化石材料。形态比较表明,这两个人类头骨化石的脑颅虽然在眶上圆枕上沟的发育程度、眶后收缩的程度、额骨横向隆起的程度、角圆枕和乳后突的发育与否、顶骨形状以及骨壁厚度的表现上有所差异,但有更多的形态性状显示出相近。这些相近表现在脑颅的长、宽、高值上;颅容量上;脑颅的低矮性上;脑颅最大宽之位置上;额骨、顶骨、枕骨之矢弧值的比例上;眶上圆枕的纤细上;顶骨的大小和矢向扁平性上;颞线位置和颞鳞顶缘的形状上;枕鳞的低宽形状上;上枕鳞与下枕鳞之间的转折形状和比例上;枕骨圆枕和枕骨圆枕上沟的发育程度上等。这两具头骨的面颅虽然有同属突颌型的面角、皆发育有鼻骨间嵴、两鼻骨组成的上部宽度与下部宽度皆差别很大,但有更多的形态性状显示出差别。这些差别表现在面型上、颜面上部扁平度上、眶形和眶型上、上颌额突外侧面的朝向上、鼻骨横向隆起程度上、鼻梁外突程度上、鼻型上、颧骨下缘外展程度上、颊高上、颧上颌下缘的形状上、上颌颧突基部的位置上以及颧结节的位置上等。因此,南京1号头骨与KNM-ER 3733头骨之间在脑颅上显示出较多的相近性状,在面颅上则显示出较多的相异性状。脑颅方面的相近性状大多具有分类上的鉴别价值。这两个头骨脑颅形态的相近支持把KNM-ER 3733头骨鉴定为"直立人"的观点;也提示了南京1号头骨的脑颅似乎保持着1百多万年前的"祖先"形态。如果直立人的某些成员在至少1百万年期间保持着形态稳定的话,则这种形态上的稳定主要是表现在脑颅形态上。这两具头骨的面颅形态上较大差异的意义,目前尚不清楚。