Cranial vault shape in fossil hominids: Fourier descriptors in norma lateralis

Two major views of human evolution have elicited considerable controversy. These are: [1] the “out of Africa” hypothesis and [2] the “multiregional” hypothesis. This paper is an attempt to try to reconcile these two scenarios using hominid cranial vault data. Elliptical Fourier functions (EFFs) were used to describe, in visual and numerical terms, the shape of the human cranial vault in norma lateralis.Using jpeg images, contours of the cranial vault of a large sample of hominid specimens were pre-processed in Photoshop CS and rotated in 2D space (positional-orientation) so that a line drawn from nasion to porion was horizontal. The cranial vault image was then digitized with 72 closely-spaced points and submitted to a specially written routine that computed EFFs normalized by scaling (size-standardization). This ensured that the representation was invariant with respect to starting point, size and orientation.Statistically significant differences were found between the H. sapiens sample and both the H. erectus and H. neanderthalensis samples. In contrast, there were no statistically significant differences between the H. erectus and H. neanderthalensis groups, leading to three conclusions: [1] the similarity in cranial vault shape between H. erectus and H. neanderthalensis suggests a single gradually evolving lineage; [2] The taxon H. heidelbergensis can be embedded into the H. erectus → H. neanderthalensis line; and [3] H. sapiens seems to be a separate evolutionary development and is considered here either as a separate species or as a possible example of an allopatric semispecies (Grant, 1977). The results here suggest that human evolution over the last 2 Ma may turn out to be neither totally multiregional or simply out of Africa but rather represents a considerably more complicated picture.     

Cranial growth in Homo erectus: how credible are the Ngandong juveniles?

Confusion exists regarding the developmental ages of numerous Asian and southeast Asian Homo erectus fossils because of Weidenreich's contention that Pithecanthropus fused its sutures prematurely relative to H. sapiens. I reevaluate the cranial developmental ages of the Ngandong “juveniles” (2, 5, 8, 9) based on a series of indicators of youth (superstructure development, suture development/fusion, and cranial thickness) and cranial contours. The Ngandong juveniles are compared with H. sapiens adults (n = 281) and subadults (n = 81) and with Ngandong and other H. erectus adults (n = 20) and subadults (n = 4). Cranial contours are assessed using bivariate plots of arc vs. chord measurements. All indicators suggest that Ngandong 5 and 9 are adults, whereas Ngandong 8 is an older juvenile or young adult and Ngandong 2 is a juvenile with a developmental age range of greater than 6 and less than 11 years. In addition, adult cranial contours and the pattern of contour development are similar between Ngandong adults and other H. erectus adults. There is nothing in the cranial contour data to suggest that Ngandong is, despite a relatively large brain, transitional in vault shape between H. erectus and H. sapiens. Am J Phys Anthropol 108:223–236, 1999. © 1999 Wiley-Liss, Inc.     

A comparative study of frontal bone morphology among Pleistocene hominin fossil groups

Features of the frontal bone that are conventionally used to distinguish among fossil hominin groups were quantitatively examined. Fifty-five fossil crania dating from the early to the late Pleistocene were analyzed. Using a modified pantograph, outlines of the frontal bone were collected along the midsagittal and two parasagittal planes. The profile from nasion to bregma, as well as two profiles above the medial and lateral sections of the orbit, respectively, extending from the orbital margin to the coronal suture were traced. The outlines were measured using Elliptical Fourier Function Analysis (EFFA), which enabled a quantification of aspects of the frontal bone that have historically been described primarily in nonmetric or linear terms. Four measurements were obtained: 1) overall morphology as expressed in the Fourier harmonic amplitudes; 2) maximum projection of the supraorbital torus at three points along the browridge (glabella and the medial and lateral aspects of the torus above the orbit); 3) maximum distance of the frontal squama from the frontal chord, capturing forehead curvature; and 4) nasion-bregma chord length. The results indicate that the midsagittal profile is significantly different among all Pleistocene groups in analyses that include both size and shape, as well as size-adjusted data. Homo erectus is significantly different from the late Pleistocene groups (Neandertals and early modern H. sapiens) in glabellar projection. Anatomically modern humans are significantly different from all other groups in both raw and size-standardized analyses of all three outlines that captured overall morphology, as well as forehead curvature and lateral supraorbital torus prominence, and middle Pleistocene Homo are significantly different in both medial and lateral overall parasagittal form. However, for the majority of analyses there were no significant differences among the Pleistocene archaic groups in supraorbital torus projection, frontal squama curvature, nasion-bregma chord length, or overall frontal bone morphology.     

A newHomo erectus cranium from Sangiran, Java

A newHomo erectus cranium was found on May 18, 1993 by Budi, a local farmer, at Sangiran. It dates from the Middle Pucangan Formation approximately 1.6–1.8 mya. The braincase is essentially complete and as is most of the face. The vault has the typicalH. erectus gable shape. There is a clear sagittal ridge beginning below the middle of the frontal squama and running to mid-parietal. Parasagittal ridges are rounded angulations halfway up the parietals, and coincide with poorly marked temporal lines. In all measurements, this skull is longer and consistently narrower than Trinil. It is chronologically and morphologically similar to the famousH. erectus skull from east Africa, KNMER-3733. Although existing much older, this new specimen is what one would expect a female counterpart to Sangiran 17 to look like.     

Allometric patterns of cranial bone thickness in fossil hominids

The interspecific allometry of five measures of total cranial bone thickness is examined in 10 extant catarrhine genera and two fossil hominid samples representing A. africanus and Asian H. erectus. Analysis of the modern sample shows that most interspecific variation in vault thickness can be accounted for by variation in body size. Correlation values are moderate to high (r = 0.75–0.98), and all variables exhibit positive allometry. The bone thickness:body mass relationship of modern humans broadly conforms with that of other primates. However, in the distribution of relative thickness throughout the skull, H. sapiens is distinguished by relative thickening of the parietal and extreme relative thinning of the temporal squama. The bone thickness:body mass relationship in the two early hominid species is examined using published mean body weight estimates generated from post-cranial predictor variables. A. africanus exhibits great similarity to modern humans in its relation to the catarrhine regression data and in the distribution of relative thickness throughout the skull. H. erectus also shows a modern human-like pattern in the distribution of its relative thickness; however, its bone thickness:body mass relationship is dissimilar to that displayed by all other taxa, including the other hominid species. On the basis of these results, it is suggested that the published body weight estimate assigned to H. erectus greatly underestimates actual mean body size for Asian members of this species. © 1996 Wiley-Liss, Inc.     

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.     

Functional implications of squamosal suture size in paranthropus boisei

It has been hypothesized that the extensively overlapping temporal and parietal bones of the squamosal sutures in Paranthropus boisei are adaptations for withstanding loads associated with feeding. Finite element analysis (FEA) was used to investigate the biomechanical effects of suture size (i.e., the area of overlap between the temporal and parietal bones) on stress, strain energy, and strain ratio in the squamosal sutures of Pan troglodytes and P. boisei (specimen OH 5) during biting. Finite element models (FEMs) of OH 5 and a P. troglodytes cranium were constructed from CT scans. These models contain sutures that approximate the actual suture sizes preserved in both crania. The FEM of Pan was then modified to create two additional FEMs with squamosal sutures that are 50% smaller and 25% larger than those in the original model. Comparisons among the models test the effect of suture size on the structural integrity of the squamosal suture as the temporal squama and parietal bone move relative to each other during simulated premolar biting. Results indicate that with increasing suture size there is a decreased risk of suture failure, and that maximum stress values in the OH 5 suture were favorable compared to values in the Pan model with the normal suture size. Strain ratios suggest that shear is an important strain regime in the squamosal suture. This study is consistent with the hypothesis that larger sutures help reduce the likelihood of suture failure under high biting loads. Am J Phys Anthropol 153:260–268, 2014. © 2013 Wiley Periodicals, Inc.     

Evolution of the zygomasseteric construction in Rodentia,as revealed by a geometric morphometric analysis of the mandible of Graphiurus (Rodentia,Gliridae)

Graphiurus is a peculiar taxon among the monophyletic Gliridae (order Rodentia) in showing hystricomorphy of the zygomasseteric architecture of the skull [large infraorbital foramen (IOF), and correlative muscular arrangements). We analysed 34 extant genera taken from two groups of sciurognath rodents that share a large IOF (hystricomorph and myomorph) using elliptical Fourier transform in order to appraise whether this feature of cranial morphology was also accompanied by similar changes in mandible shape. The mandible of Graphiurus is distinct from those of all other hystricomorph sciurognath rodents in showing a more elongated coronoid process and a shorter angular process. Thus, two distinct zygomasseteric organizations (i.e. myomorphy and hystricomorphy of graphiurines) are associated with a similar mandible shape characterized by a well‐developed coronoid process. Results show that hystricomorphy of graphiurines was achieved convergently with other hystricomorph rodents. Protrogomorphy is the plesiomorphic condition in Gliridae and hystricomorphy is an autapomorphic feature of Graphiurus. © 2008 The Linnean Society of London, Zoological Journal of the Linnean Society, 2008, 154 , 807–821.     

Interspecific variation of ontogeny and skull shape among porpoises (Phocoenidae)

All extant members of Phocoenidae (porpoises) have been characterized as pedomorphic based on skeletal characters. To investigate the ontogenetic background for pedomorphosis and assess interspecific differences in ontogeny among phocoenids, samples of the six extant species were compared in terms of development of both epiphyseal and cranial suture fusion. Across all species, full maturity of the vertebral column was rare. Vertebral epiphyseal development did not progress so far in most Phocoena phocoena as in Phocoenoides dalli and Phocoena dioptrica. P. phocoena, Phocoena spinipinnis, Ph. dalli, and P. dioptrica, for which large series were available, were further compared in terms of ontogeny of cranial shape by three‐dimensional geometric morphometrics. Ph. dalli and P. dioptrica generally showed further development of cranial sutures than the other species. Postnatal skull shape development was similar for all species studied; the majority of interspecific shape differences are present at parturition. Smaller species had a higher rate of shape development relative to growth in size than Ph. dalli and P. dioptrica, but they still showed less allometric development due to less postnatal growth. Interspecific shape differences indicate phylogenetic relationships similar to that proposed based on morphology or convergent evolution of the two pelagic species, Ph. dalli and P. dioptrica, under the scenarios suggested by recent molecular studies. A shape trend coinciding with habitat preference was detected; in species with pelagic preference the position and orientation of the foramen magnum aligned the skull with the vertebral column; the rostrum showed less ventral inclination, and the facial region was larger and more concave in lateral aspect. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.     

Size and shape of human cranial sutures--a new scoring method

A method for the differentiation of sutural patterns of the human cranial vault is introduced. Three criteria of differentiation are considered, one for size and two for shape: 1) maximal shape extension; 2) basic configuration; 3) secondary protrusion. The method is illustrated here for the coronal and lambdoid sutures of 70 recent Italian skulls (35 adult males and 35 adult females). Differences between coronal and lambdoid sutural size and shape can be detected analytically; for example, the coronal suture commonly shows lesser degrees of shape extension, a simpler basic configuration, and an absence of secondary protrusion. Heterogeneity within each suture, as well as a relationship among corresponding sections and between the three criteria adopted, have been also observed; symmetry predominates for both the sutures, and sexual differences are slight.     

A SINGLE LINEAGE IN EARLY PLEISTOCENE HOMO: SIZE VARIATION CONTINUITY IN EARLY PLEISTOCENE HOMO CRANIA FROM EAST AFRICA AND GEORGIA

The relationship between Homo habilis and early African Homo erectus has been contentious because H. habilis was hypothesized to be an evolutionary stage between Australopithecus and H. erectus, more than a half‐century ago. Recent work re‐dating key African early Homo localities and the discovery of new fossils in East Africa and Georgia provide the opportunity for a productive re‐evaluation of this topic. Here, we test the hypothesis that the cranial sample from East Africa and Georgia represents a single evolutionary lineage of Homo spanning the approximately 1.9–1.5 Mya time period, consisting of specimens attributed to H. habilis and H. erectus. To address issues of small sample sizes in each time period, and uneven representation of cranial data, we developed a novel nonparametric randomization technique based on the variance in an index of pairwise difference from a broad set of fossil comparisons. We fail to reject the hypothesis of a single lineage this period by identifying a strong, time‐dependent pattern of variation throughout the sequence. These results suggest the need for a reappraisal of fossil evidence from other regions within this time period and highlight the critical nature of the Plio‐Pleistocene boundary for understanding the early evolution of the genus Homo.     

Enlargement of the temporalis muscle and alterations in the lateral cranial vault

The purpose of this study was to test the hypothesis that increasedmasticatory muscle accompanied morphologic changes in the temporalbone and squamosal suture. Ten mice deficient for the proteinmyostatin (Mstn –/–) had significantly increasedskeletal muscle mass and were compared with nine controls (Mstn+/+). Variables measured include linear and areal metrics describingtemporal size and temporal bone shape as well as the extentof the area of the squamosal suture that overlaps, or bevels,with parietal bones. Mstn–/– mice showed significantlylarger temporalis muscles. Their temporal bones showed significantlydecreased size as well as decreased beveling of the squamosalsuture. These decreases were absolute as well as relative andwere not restricted to either vertical or horizontal axes. Theincreased masticatory musculature of Myostatin-null mice hada shrinking effect on the temporal aspect of the cranium. Theseresults are inconsistent with the interpretation that increasedtemporalis mass induces morphologic changes in temporal bonethat compensate for putative increases in compressive forcestransduced at this region. Rather than increase in the areaof overlap between two calvarial bones, potential increase inbiomechanical loading along the temporal squama led to a smallerbevel which would presumably weaken this joint. It is unclearwhy this is so. Either compressive forces are not anabolic tosuture beveling or they do upregulate growth of the suture bevel,with compression not being the primary loading regime at thissuture.     

Functional implications of dicynodont cranial suture morphology

Cranial suture morphology of Lystrosaurus and the generalized dicynodont Oudenodon was investigated to determine the strain environment during mastication, which in turn may indicate a difference in cranial function between the two taxa. Finite element (FE) analysis indicated that less strain accumulated in the cranium of Lystrosaurus during orthal bite simulations than in Oudenodon. Despite the overall difference in strain magnitude, moderate to high FE‐predicted strain accumulated in similar areas of the cranium of both taxa. The suture morphology in these cranial regions of Lystrosaurus and Oudenodon was investigated further by examination of histological sections and supplemented by observations of serial sections and computed tomography (CT) scans. The predominant type of strain from selected blocks of finite elements that contain sutures was determined, enabling comparison of suture morphology to strain type. Drawing from strain‐suture correlations established in extant taxa, the observed patterns of sutural morphology for both dicynodonts were used to deduce cranial function. The moderate to high compressive and tensile strain experienced by the infraorbital bar, zygomatic arch, and postorbital bar of Oudenodon and Lystrosaurus may have been decreased by small adjustive movements at the scarf sutures in those regions. Disparities in cranial suture morphology between the two taxa may reflect differences in cranial function. For instance, the tongue and groove morphology of the postorbital‐parietal suture in Oudenodon could have withstood the higher FE‐predicted tensile strain in the posterior skull roof. The scarf premaxilla‐nasal suture of Lystrosaurus provided an additional region of sutural mobility in the anterior surface of the snout, suggesting that Lystrosaurus may have employed a different biting regime than Oudenodon. The morphology of several sutures sampled in this study correlated with the FE‐predicted strain, although other cranial functional hypotheses remain to be tested. J. Morphol., 2010. © 2010 Wiley‐Liss, Inc.     

Geometric variation of the frontal squama in the genus homo: Frontal bulging and the origin of modern human morphology

The majority of studies of frontal bone morphology in paleoanthropology have analyzed the frontal squama and the browridge as a single unit, mixing information from different functional elements. Taking into account that the bulging of the frontal bone is often described as a species‐specific trait of Homo sapiens, in this article we analyze variation in the midsagittal profile of the genus Homo, focusing on the frontal squama alone, using landmark‐based superimpositions and principal components analysis. Our results demonstrate that anatomically modern humans are definitely separated from extinct human taxa on the basis of frontal bulging. However, there is minor overlap among these groups, indicating that it is necessary to exercise caution when using this trait alone to make taxonomic inferences on individual specimens. Early modern humans do not show differences with recent modern humans, and “transitional” individuals such as Jebel Irhoud 1, Maba, and Florisbad, show modern‐like frontal squama morphology. The bulging of the frontal squama in modern humans may represent a structural consequence of more general cranial changes, or it could be a response to changes in the morphology of the underlying prefrontal brain elements. A subtle difference between Neandertals and the Afro‐European Middle Pleistocene Homo sample is associated with flattening at bregma in the former group, a result that merits further investigation. Am J Phys Anthropol, 2013. © 2013 Wiley Periodicals, Inc     

Ontogenetic shape changes in Pomacentridae (Teleostei,Perciformes) and their relationships with feeding strategies: a geometric morphometric approach

The present study explores the shape changes of cranial structures directly involved in food capturing during growth after reef settlement in two species of Pomacentridae (Dascyllus aruanus and Pomacentrus pavo). Landmark‐based geometric morphometrics were used to study allometric patterns and related shape changes in four skeletal units: neurocranium, suspensorium and opercle, mandible and premaxilla. At settlement, the larvae of both species have a relatively similar morphology, especially with respect to the mandible. Their shapes suggest a feeding mode defined as ram/suction‐feeding. Ontogenetic shape changes show a shift to a suction feeding mode of prey capture. The main transformations involved are an increase in height of the suspensorium and the opercle, an elevation of the supraoccipital crest, a relative shortening of the mandible, and a lengthening of the ascending process of the premaxilla. Shape changes of the mandible in the two studied species also reflect an increase of biting capacities. The high disparity between adult shape results from differences in the rate and in the length of ontogenetic trajectories, from divergence of the ontogenetic trajectories (neurocranium, mandible, and premaxilla) and parallel shifts of the trajectories in the size‐shape space (suspensorium and opercle). In an evolutionary context, allometric heterochronies during ontogeny of different skeletal unit of the head may be considered as a basis for the explanation of the diversity of damselfishes. © 2008 The Linnean Society of London, Biological Journal of the Linnean Society, 2008, 95 , 92–105.     

Occipital bunning among later pleistocene hominids

Occipital bunning is a posterior projection of the occipital squama, which occurs in varying frequencies in samples of archaic Homo sapiens, Upper Pleistocene anatomically modern humans, and recent humans. It can be best interpreted as a product of the timing of posterior cerebral growth relative to the growth of the cranial vault bones. It is not a feature that was unique to the Neandertals.