Avian skull morphological evolution: exploring exo- and endocranial covariation with two-block partial least squares

While rostral variation has been the subject of detailed avian evolutionary research, avian skull organization, characterized by a flexed or extended appearance of the skull, has eventually become neglected by mainstream evolutionary inquiries. This study aims to recapture its significance, evaluating possible functional, phylogenetic and developmental factors that may be underlying it. In order to estimate which, and how, elements of the skull intervene in patterning the skull we tested the statistical interplay between a series of old mid-sagittal angular measurements (mostly endocranial) in combination with newly obtained skull metrics based on landmark superimposition methods (exclusively exocranial shape), by means of the statistic-morphometric technique of two-block partial least squares. As classic literature anticipated, we found that the external appearance of the skull corresponds to the way in which the plane of the caudal cranial base is oriented, in connection with the orientations of the plane of the foramen magnum and of the lateral semicircular canal. The pattern of covariation found between metrics conveys flexed or extended appearances of the skull implicitly within a single and statistically significant dimension of covariation. Marked shape changes with which angles covary concentrate at the supraoccipital bone, the cranial base and the antorbital window, whereas the plane measuring the orientation of the anterior portion of the rostrum does not intervene. Statistical covariance between elements of the caudal cranial base and the occiput inplies that morphological integration underlies avian skull macroevolutionary organization as a by-product of the regional concordance of such correlated elements within the early embryonic chordal domain of mesodermic origin.     

以乳突切迹和翼钩为基点的侧颅底分区方法

目的:探讨以乳突切迹和翼钩为基点的侧颅底分区新方法.方法:在乳突切迹后缘、翼钩、枕骨大孔前缘中点和颧根四个结构间相互连线,区分侧颅底并测量连线的长度.结果:乳突切迹与侧颅底重要结构的关系密切,切迹后缘与翼钩连线和正中线将侧颅底分成内、外侧两个大的三角区,每个区再分成前后两个三角区共四个三角区,即腭和颞下三角、咽三角、关节和听三角、血管神经三角,其中血管神经三角的三边长度左右侧分别为(74.52±5.47)mm和(74.66±5.41)mm、(59.77±3.84)mm和(59.67±3.56)mm、(42.23±3.11)mm和(42.48±2.60)mm.结论:本研究提供了新的侧颅底分区方法,且血管神经三角的区域划分更为科学,为临床侧颅底手术入路和定位提供了解剖学参考.     

Brief communication: foramen magnum-carotid foramina relationship: is it useful for species designation?

The anterior placement of the foramen magnum is often used to indicate bipedalism and therefore to distinguish hominid from nonhominid fossils. Often, only fragmentary cranial remains are found, and the placement of the foramen magnum must be determined by its relationship to another landmark. The purpose of this study was to test if hominid crania could be distinguished from nonhominid crania based on the relationship between the foramen magnum and the carotid foramina, and therefore to determine if the carotid foramina can be used to determine the anteriorness of the foramen magnum. The samples consisted of 16 modern human crania and 19 modern chimpanzee (Pan troglodytes) crania. Linear measurements were taken of (1) the distance from the anterior border of the foramen magnum to a chord connecting the carotid foramina and (2) total cranial length. An index of the distance of the foramen magnum from the bicarotid chord as a proportion of total cranial length was calculated to control for differences due to body size. Results indicate that on average, human crania can be distinguished from chimpanzee crania by using either (1) the distance of the foramen magnum from the bicarotid chord as a linear measurement or (2) this linear measurement as a proportion of total cranial length. Both measures are significantly smaller in the human sample; however, there was considerable overlap between species, indicating that the distance of the foramen magnum from the bicarotid chord is not a certain indicator for individual specimens.     

乳突切迹与侧颅底重要结构的解剖学观测及应用

目的：通过测量乳突切迹与侧颅底重要骨性结构的距离,为临床相关应用提供解剖学参考。方法：取成人颅骨标本50例（去颅盖标本8例,整颅42例）100侧,测量乳突切迹及其与侧颅底重要孔、裂和管的距离。结果：左右侧乳突切迹后缘距茎乳孔、颈静脉孔外侧缘、颈动脉管外口后缘、破裂孔、棘孔、卵圆孔距离分别为25.16±3.73cm和25.02±3.58cm、30.92±3.50cm和30.45±3.49cm、38.22±3.57cm和38.14±3.43cm、57.23±3.71cm和57.14±3.44cm、47.94±3.83cm和48.32±3.54cm、53.70±3.98cm和53.55±3.75cm。结论：以乳突切迹后缘做为侧颅底手术的定位标志能够为临床相关应用提供较方便、准确的定位参考。     

颧根与侧颅底重要结构的解剖学测量

目的：通过测量颧根与侧颅底各重要结构间的距离关系,为临床侧颅底外科手术治疗提供定位参考。方法：取成人颅骨标本50例（去颅盖标本8例,整颅42例）100侧,用游标卡尺、圆规和直尺测量颧根与侧颅底重要结构的距离。结果：实验测得左右侧颧根与外耳门前缘中点、乳突尖、茎突、翼突外侧板根部、舌下神经管外口、茎乳孔、颈静脉孔外缘、颈动脉管外口后缘、棘孔、卵圆孔、破裂孔的距离分别为22.30±2.84mm和22.02±3.27mm、40.37±3.21mm和40.56±3.54mm、32.53±2.78mm和32.92±2.68mm、35.13±3.14mm和35.19±2.74mm、49.29±2.88mm和48.98±2.87mm、32.92±2.44mm和33.05±2.61mm、35.15±2.86mm和34.68±3.13mm、33.17±2.78mm和33.17±2.72mm、28.83±2.62mm和28.68±2.63mm、31.15±2.76mm和31.49±2.73mm、43.67±3.32mm和44.15±3.02mm,左右侧数据无差异（P〉0.05）。结论：颧根可以作为侧颅底外科手术的定位标志,为直视条件下经颞下等入路的侧颅底外科手术提供解剖学依据。     

Position and orientation of the foramen magnum in higher primates

The location of the foramen magnum, with respect to the longitudinal axis of the cranium, and its orientation with respect to the Frankfurt Horizontal, have been studied in a total of 328 modern human and Pan crania. The samples were chosen in order to examine the effect of overall size difference on foramen magnum disposition. Foramen position (expressed as three indices) and inclination are relatively invariant among the modern human samples, but the foramen magnum is consistently, and statistically significantly, more anteriorly located in Pan paniscus than in Pan troglodytes. Sexual dimorphism is virtually non-existent. There is an apparent allometric effect on foramen position, but not on inclination, so that larger crania in the modern human and Pan paniscus samples tend to have more posteriorly situated foramina. The disposition of the foramen is unrelated to cranial base angle or facial prognathism, except that in Pan paniscus its relative anterior location is linked with the more flexed cranial base in that species. These results provide a comparative context for the examination of differences in foramen magnum disposition in fossil hominids. Differences in foramen magnum position and orientation between KNM-ER 1813 and A. africanus are most unlikely to be due to within-taxon variability.     

The cranial base of Australopithecus afarensis: new insights from the female skull

Cranial base morphology differs among hominoids in ways that are usually attributed to some combination of an enlarged brain, retracted face and upright locomotion in humans. The human foramen magnum is anteriorly inclined and, with the occipital condyles, is forwardly located on a broad, short and flexed basicranium; the petrous elements are coronally rotated; the glenoid region is topographically complex; the nuchal lines are low; and the nuchal plane is horizontal. Australopithecus afarensis (3.7–3.0 Ma) is the earliest known species of the australopith grade in which the adult cranial base can be assessed comprehensively. This region of the adult skull was known from fragments in the 1970s, but renewed fieldwork beginning in the 1990s at the Hadar site, Ethiopia (3.4–3.0 Ma), recovered two nearly complete crania and major portions of a third, each associated with a mandible. These new specimens confirm that in small-brained, bipedal Australopithecus the foramen magnum and occipital condyles were anteriorly sited, as in humans, but without the foramen''s forward inclination. In the large male A.L. 444-2 this is associated with a short basal axis, a bilateral expansion of the base, and an inferiorly rotated, flexed occipital squama—all derived characters shared by later australopiths and humans. However, in A.L. 822-1 (a female) a more primitive morphology is present: although the foramen and condyles reside anteriorly on a short base, the nuchal lines are very high, the nuchal plane is very steep, and the base is as relatively narrow centrally. A.L. 822-1 illuminates fragmentary specimens in the 1970s Hadar collection that hint at aspects of this primitive suite, suggesting that it is a common pattern in the A. afarensis hypodigm. We explore the implications of these specimens for sexual dimorphism and evolutionary scenarios of functional integration in the hominin cranial base.     

Correlation coefficients in human skulls: significant sexual differences

This study of 500 male and female skulls proves: significant differences of correlation coefficients (p less than or equal to 0.01) occur between the two sexes. The combined variables are: 1. bizygomatic breadth (45)/skull base length (5) 2. bizygomatic breadth (45)/foramen magnum length (7) 3. bizygomatic breadth (45)/foramen magnum breadth (16) 4. bizygomatic breadth (45)/basion-bregma height (17) 5. foramen magnum breadth (16)/skull base length (5) Female skulls have generally higher correlation coefficients values than males which is interpreted as an indication of homogeneous growth.     

Comparative myology of the hominoid cranial base. I. The muscular relationships and bony attachments of the digastric muscle

This paper aims to document accurately the soft tissue anatomy and bony attachments of the posterior belly of the digastric muscle and other closely related muscles in the mastoid region of extant hominoids and fossil hominids. Five wet specimens including individuals of Pan, Gorilla and Pongo were dissected and described. Eight casts of fossil hominid cranial bases were also studied along with measurements and notes made from the same original fossil hominid specimens to assess their soft tissue markings in the light of the findings for the three great apes. The results indicate that whereas the attachment of the posterior belly of the digastric muscle in Homo sapiens is associated with a deep groove or fossa, it originates from a widened area and leaves no bony markings on the cranial base of the three great apes. Following a change in the position of the foramen magnum and the occipital condyles in hominids and H. sapiens the insertion of the posterior belly of the digastric has remained posteriorly positioned but has become compressed into a deep groove. It is likely that this has come about by the displacement of the more medial soft tissue structures which have been moved laterally away from the occipital condyles.     

Measurements on the grade of the human skull base and angle determinations]

1. The angle between the Os sphenoidale and the Clivus has been determined on 106 skulls. The mean was 117.68 degrees. 2. The altitude of the Foramen magnum with respect to the Frankfort plane was determined at -1.96 cm on the average. 3. The angle of inclination of the Foramen magnum to the Frankfort plane was determined from the altitude of the Basion and Opisthion and from the length of the Foramen magnum. The mean was -13.71 degrees. 4. In the inner base of the skull the altitudes of following structures have been determined: Planum sphenoidale (mean 2.34 cm), ground of the Fossa hypophysialis (1.32 cm), ground of the middle cranial fossa (-0.03 cm), ground of the posterior cranial fossa (-2.30 cm), upper edge of the Os petrosum (1.12 cm), Tuberculum jugulare (-0.21 cm), ground of the Meatus acusticus internus (0.45 cm). 5. In order to provide numerical data for the step-like building of the inner base of the skull, the differences between the altitudes of important structures from the inner base have been determined: Margo superior partis petrosae - Tuberculum jugulare (mean 1.35 cm). Opisthion - ground of the posterior cranial fossa (0.11 cm), Planum sphenoidale - ground of the middle cranial fossa (2.38 cm), ground of the middle cranial fossa - ground of the posterior cranial fossa (2.27 cm), Margo superior partis petrosae - ground of the posterior cranial fossa (3.35 cm), Margo superior partis petrosae - ground of the middle cranial fossa (1.15 cm).     

Brain size and the human cranial base: a prenatal perspective

Pivotally positioned as the interface between the neurocranium and the face, the cranial base has long been recognized as a key area to our understanding of the origins of modern human skull form. Compared with other primates, modern humans have more coronally orientated petrous bones and a higher degree of basicranial flexion, resulting in a deeper and wider posterior cranial fossa. It has been argued that this derived condition results from a phylogenetic increase in the size of the brain and its subcomponents (infra- and supratentorial volumes) relative to corresponding lengths of the cranial base (posterior and anterior, respectively). The purpose of this study was to test such evolutionary hypotheses in a prenatal ontogenetic context. We measured the degree of basicranial flexion, petrous reorientation, base lengths, and endocranial volumes from high-resolution magnetic resonance images (hrMRI) of 46 human fetuses ranging from 10-29 weeks of gestation. Bivariate comparisons with age revealed a number of temporal trends during the period investigated, most notable of which were coronal rotation of the petrous bones and basicranial retroflexion (flattening). Importantly, the results reveal significant increases of relative endocranial sizes across the sample, and the hypotheses therefore predict correlated variations of cranial base flexion and petrous orientation in accordance with these increases. Statistical analyses did not yield results as predicted by the hypotheses. Thus, the propositions that base flexion and petrous reorientation are due to increases of relative endocranial sizes were not corroborated by the findings of this study, at least for the period investigated.     

以乳突切迹和翼钩为基点的侧颅底分区方法

目的：探讨以乳突切迹和翼钩为基点的侧颅底分区新方法。方法：在乳突切迹后缘、翼钩、枕骨大孔前缘中点和颧根四个结构间相互连线,区分侧颅底并测量连线的长度。结果：乳突切迹与侧颅底重要结构的关系密切,切迹后缘与翼钩连线和正中线将侧颅底分成内、外侧两个大的三角区,每个区再分成前后两个三角区共四个三角区,即腭和颞下三角、咽三角、关节和听三角、血管神经三角,其中血管神经三角的三边长度左右侧分别为（74.52±5.47）mm和（74.66±5.41）mm、（59.77±3.84）mm和（59.67±3.56）mm、（42.23±3.11）mm和（42.48±2.60）mm。结论：本研究提供了新的侧颅底分区方法,且血管神经三角的区域划分更为科学,为临床侧颅底手术入路和定位提供了解剖学参考。     

Palaeoanthropological discoveries in Indonesia with special reference to the finds of the last two decades

This article reviews palaeoanthropological research in Indonesia since 1889 in terms of fossil discoveries. Of the three periods identified, the second one (1931–1941) resulted in more finds than the other two. Most finds are skull fragments of Pithecanthropus erectus, from the Middle Pleistocene Kabuh formation, and from the site of Sangiran. New code numbers are given to the finds to facilitate discussion, especially in the cases of previous misnumbering and controversial specimens.K/Ar dating gives an antiquity of 1·9 ± 0·4 million years for the Jetis beds at Perning (site of the Mojokerto juvenile calvaria) and 830,000 years for the Trinil beds at Sangiran (sites of Sangiran 10 and 12 calottes). Work on chronometric dating is continuing.Several re-examinations of earlier finds from Java have been performed in the last two decades. Finds during this period are reviewed and described, comprising one calvaria, two skull caps, two mandibular fragments, skull vault and base fragments, and teeth. One mandibular fragment is related to Meganthropus, but no skull fragments are associated with this to make its taxonomic status clearer. For the first time portion of the cranial base anterior to the foramen magnum and the zygomatic bone of Pithecanthropus erectus are available.The fossil finds demonstrate the genetic consistency of Pithecanthropus as revealed by the morphology of the skull. Palaeoanthropological research goes on, slowly but surely, especially in the Sangiran dome area of Central Java.     

Major cranial changes during Triceratops ontogeny

This is the first cranial ontogenetic assessment of Triceratops, the well-known Late Cretaceous dinosaur distinguished by three horns and a massive parietal-squamosal frill. Our analysis is based on a growth series of 10 skulls, ranging from a 38 cm long baby skull to about 2 m long adult skulls. Four growth stages correspond to a suite of ontogenetic characters expressed in the postorbital horns, frill, nasal, epinasal horn and epoccipitals. Postorbital horns are straight stubs in early ontogeny, curve posteriorly in juveniles, straighten in subadults and recurve anteriorly in adults. The posterior margin of the baby frill is deeply scalloped. In early juveniles, the frill margin becomes ornamented by 17-19 delta-shaped epoccipitals. Epoccipitals are dorsoventrally compressed in subadults, strongly compressed and elongated in adults and ultimately merge onto the posterior frill margin in older adults. Ontogenetic trends within and between growth stages include: posterior frill margin transitions from scalloped to wavy and smooth; progressive exclusion of the supraoccipital from the foramen magnum; internal hollowing at the base of the postorbital horns; closure of the midline nasal suture; fusion of the epinasal onto the nasals; and epinasal expansion into a morphologically variable nasal horn. We hypothesize that the changes in horn orientation and epoccipital shape function to allow visual identity of juveniles, and signal their attainment of sexual maturity.     

Brief communication: Sexual dimorphism of the juvenile basicranium

The purpose of this article is to examine the level of sexual dimorphism exhibited in the foramen magnum and occipital condyles of juveniles, and to test the utility of this sexual dimorphism for estimating sex. Using five basicranial measurements taken from 36 juveniles of known sex and age from the Lisbon documented collection (Portugal), we evaluated sexual dimorphism in the juvenile cranial base. Our application of a method previously applied solely to adults indicated that the basicranium is sexually dimorphic in juveniles, with larger foramen magnum and occipital condyle dimensions observed in males. Significant univariate differences between males and females were found for length and breadth of the foramen magnum, and breadth of the left occipital condyle. Using these measurements, multivariate discriminant analysis indicated that sex was correctly assigned 75.8% of time. Obtained accuracy, however, was lower than reported by previous studies of adult samples. We suggest that this discrepancy is a result of population variability rather than age. Am J Phys Anthropol 2010. © 2009 Wiley‐Liss, Inc.     

The role of basicranial synchondroses in flexure processes and ontogenetic development of the skull base

The contribution of the basicranial synchondroses in the growth of neurocranial length and ontogenetic development of the cranial base were investigated. The study concentrated on the midsphenoidal synchondrosis and its delayed fusion in nonhuman primates when compared to man, and on the spheno-occipital synchondrosis. The mode and time of fusion of both growth centers were observed, and their role in the ontogenetic growth changes (flattening processes) of the cranial base were established. The chondrogenic ossification of midsphenoidal and spheno-occipital synchondroses was studied on 20 skulls of Macaca mulatta females, ranging in age from newborn specimens to those 24 months old. The technique of in vivo tetracycline bone labeling was used for histologic evaluation of the material. Different chondrogenic growth patterns were observed in both synchondroses. The endochondral activity of the spheno-occipital synchondrosis increased with age, from a nonactive narrow cartilaginous column in the neonatal specimen to a broad band with high chondrogenic ossification in the 24-month-old specimens. This growth center contributes to elongation of the posterior portion of the cranial base and is a secondary factor in its flexion. The midsphenoidal synchondrosis seems to be the primary factor in the mode of flexure of the cranial base in Macaques. This growth center is very active in the first ten months of life but later exhibits cessation of chondrogenic activity and long remains unfused. The first signs of fusion were observed as late as 72 months of age. At the same time, the continuous process of cranial base flattening showed the first signs of tapering off.     

颅底孔在多层CT三维重建中的测量研究

探讨颅底MSCT三维重建图像效果及主要孔的正常值,采用多层螺旋CT(MSCT),对200名正常成人(男100名,女100名)进行颅脑扫描,利用电子计算机三维重建程序立体地显示颅底外面的卵圆孔、棘孔、颈动脉管外口、破裂孔、茎乳孔、枕骨大孔,并观察其形态和测出其内径及其性差。     

The relationship between cranial base and maxillo-facial morphology in Egyptian children

The aim of this study was to investigate the relationship between the cranial base and maxillo-facial morphology in Egyptian children. Data were obtained from 95 lateral cephalograms for 61 boys and 34 girls aged from 7.5 to 9.5 years with mean age 8.5 years. Eighteen linear and 14 angular measurements were derived from 40 landmarks and recorded from the standardized radiographs that were traced following methods formerly defined and described. A principal component analysis of linear and angular measurements showed that anterior and posterior cranial base lengths and cranial base angle were closely associated in different ways with different aspects of maxillo-facial morphology in both sexes. This was more pronounced in boys. A significant positive relationship was found between anterior cranial base length and most of the variables describing the maxillo-facial morphology in both sexes. Posterior cranial base length was significantly correlated to the facial depth. The cranial base angle showed a significant negative correlation with the antero-posterior position of maxilla and mandible (SNA=-0.34, SNB=-0.27 and ANB=-0.24). In conclusion, cranial base configuration plays an important role in maxillo-facial morphology.     

Fibrous dysplasia of the cranial bones: a case report and review of the literature

Fibrous dysplasia (FD) is a relatively uncommon disorder that affects primarily the cranial region; its occurrence in the cranial base in combination with hindbrain herniation and aneurysmal bone cyst (ABC) constitutes an extremely rare condition. We report a case of polyostotic fibrous dysplasia with progressive occipital, temporal, and clival involvement. Clinical findings and differential diagnosis with special emphasis on the imaging features were discussed. A small posterior fossa volume has been thought to lead to hind brain herniation. The resultant obstruction to the CSF pathways at the level of the foramen magnum has been implicated in the development and subsequent progression of syringobulbia.