An ontogenetic perspective on the study of sexual dimorphism, phylogenetic variability, and allometry of the skull of European ground squirrel, Spermophilus citellus (Linnaeus, 1766)

Most studies of morphological variability in or among species are performed on adult specimens. However, it has been proven that knowledge of the patterns of size and shape changes and their covariation during ontogeny is of great value for the understanding of the processes that produce morphological variation. In this study, we investigated the patterns of sexual dimorphism, phylogenetic variability, and ontogenetic allometry in the Spermophilus citellus with geometric morphometrics applied to cross-sectional ontogenetic data of 189 skulls from three populations (originating from Burgenland, Banat, and Dojran) belonging to two phylogenetic lineages (the Northern and Southern). Our results indicate that sexual dimorphism in the ventral cranium of S. citellus is expressed only in skull size and becomes apparent just before or after the first hibernation because of accelerated growth in juvenile males. Sexes had the same pattern of ontogenetic allometry. Populations from Banat and Dojran, belonging to different phylogroups, were the most different in size but had the most similar adult skull shape. Phylogenetic relations among populations, therefore, did not reflect skull morphology, which is probably under a significant influence of ecological factors. Populations had parallel allometric trajectories, indicating that alterations in development probably occur prenatally. The species’ allometric relations during cranial growth showed characteristic nonlinear trajectories in the two northern populations, with accelerated shape changes in juveniles and continued but almost isometric growth in adults. The adult cranial shape was reached before sexual maturity of both sexes and adult size after sexual maturity. The majority of shape changes during growth are probably correlated with the shift from a liquid to a solid diet and to a lesser degree due to allometric scaling, which explained only 20 % of total shape variation. As expected, viscerocranial components grew with positive and neurocranial with negative allometry.     

Sexual size dimorphism and morphological divergence in the yellow‐whiskered Greenbul (Andropadus latirostris) in the Guineo‐Congolian rainforest of Africa

Evidence of sexual dimorphism in body size and the existence of morphological differences were studied in the yellow‐whiskered Greenbul Andropadus latirostris. We measured fresh body weight and seven linear parameters of external morphology in mature individuals of this species from three localities in Cameroon and two localities in Ghana. Based on general linear model analysis, we showed that males are significantly larger than females. We applied a discriminant analysis on eight morphometric parameters to create two discriminant functions, one for each country. The overall rate of well‐classified birds was 93.3% for Cameroon and 92.7% for Ghana. Wing length was the most accurate character for separating the sexes in both study areas. Significant sexual size dimorphism might be explained by sexual selection on male competitive ability and intraspecific competition. We also found morphological divergence in this species between the two study areas, including marked differences in size of the beak. This work provides statistical evidence of a substantial sexual size dimorphism in A. latirostris and geographic variation in morphology.     

Environment and space as drivers of variation in skull shape in two widely distributed South‐American Tayassuidae,Pecari tajacu and Tayassu pecari (Mammalia: Cetartiodactyla)

The influence of the environment on the geographical variation of morphological traits has been recognized in a number of taxa. Pecari tajacu and Tayassu pecari are ideal models to investigate intraspecific geographic variation in skull because of their wide and heterogeneous geographical distribution in South America. We used geometric morphometric procedures to examine the geographical variation in skull shape of 294 adult specimens of these species from 134 localities. We quantified to what extent skull shape variation was explained by environment, skull size and geographical space using variation partitioning analysis. We detected a strong pattern of geographic variation for P. tajacu skull shape, but not for T. pecari. The environment seems to be the major selective force that drives skull shape variation in both species. Nevertheless, other spatially structured processes (e.g. genetic drift, gene flow) might also have affected variation in the skull shape of the more widespread species P. tajacu. Allometric relationships might reflect the biomechanical constraints that are thought to be strong enough to limit size‐related changes in T. pecari skull shape.     

The relationship between cranial morphology,bite performance,diet and habitat in a radiation of dwarf chameleon (Bradypodion)

Many animals show unique morphological and behavioural adaptations to specific habitats. In particular, variation in cranial morphology is known to influence feeding performance, which in turn influences dietary habits and, ultimately, fitness. Dietary separation is an important means of partitioning ecological niches and avoiding inter‐ and intraspecific competition. Consequently, differences in dietary resources may help explain phenotypic divergence in closely‐related species occupying different habitats, as well as sexual dimorphism. We test this hypothesis on five phenotypic forms of a recent radiation of dwarf chameleons (Bradypodion) that vary extensively in habitat use and cranial morphology. By examining stomach contents, the dietary composition of each phenotypic form is compared to investigate potential differences in feeding strategies. Overall, chameleons in the present study exhibit considerable dietary overlap (at both inter‐ and intraspecific levels), indicating that diet is not a major driver of variation in cranial morphology within this radiation. However, the stomachs of closed‐canopy females were found to contain more prey items than male stomachs, possibly indicating that females require a greater caloric intake than their male counterparts.     

Ontogenetic development and sexual dimorphism of franciscana dolphin skull: A 3D geometric morphometric approach

The aim of this work was to study the postnatal ontogenetic development of Pontoporia blainvillei skull, identifying major changes on shape, and relating them to relevant factors in the life history of the species. We analyzed a complete ontogenetic series (73♂, 83♀) with three‐dimensional geometric morphometric techniques. Immature dolphins showed a very well‐developed braincase and a poorly developed rostrum, and the principal postnatal changes affected the rostrum and the temporal fossa, both structures implied functionally to the feeding apparatus, thus suggesting a specialized mode for catch fast prey in P. blainvillei. Osseous elements associated with sound production were already well developed on immature dolphins, suggesting the importance of this apparatus since the beginning of postnatal life. Sexual dimorphism was detected on both shape and size variables. Females were bigger than males, in accordance with previous studies. Shape differences between sexes were found on the posterior part of premaxillaries and external bony nares (P < 0.01), suggesting that this sexual dimorphism is related to differences on vocalization capabilities. J. Morphol. 275:1366–1375, 2014. © 2014 Wiley Periodicals, Inc.     

Analysis of body shape for differentiating among species of rajids

This study sought to differentiate the species of skates encountered in Gulf St Vincent (GSV), South Australia using normalized elliptical Fourier analysis of body shape. Significant intraspecific variation was observed among whole body shapes. This was overcome by limiting subsequent analyses to the anterior snout region, where significant differences in shape were detected among the species examined and provided a high degree of classification success for the skates of GSV. More generally, this approach has the potential to provide a cost‐ and time‐efficient means of discrimination among species of skates. Further research is required to investigate the potentially confounding effects of sexual dimorphism and ontogenetic variation in growth to improve the efficacy of the body shape analysis of the skates and batoids species in general. In addition, this approach requires considerable development to facilitate implementation in a fishery setting.     

Variation in guenon skulls (II): sexual dimorphism

Patterns of size and shape sexual dimorphism in adult guenons were examined using a large sample of skulls from almost all living species. Within species, sexual dimorphism in skull shape follows the direction of size-related shape variation of adults, is proportional to differences in size, and tends to be larger in large-bodied species. Interspecific divergence among shape trajectories, which explain within species sex differences, are small (i.e., trajectories of most species are nearly parallel). Thus, changes in relative proportions of skull regions that account for the distinctive shape of females and males are relatively conserved across species, and their magnitude largely depends on differences in size between sexes. A conservative pattern of size-related sexual dimorphism and a model of interspecific divergence in shape which strongly reflects size differences suggest a major role of size and size-related shape variation in the guenon radiation. It is possible that in the guenons, as in the neotropical primates (with whom they have obvious parallels), size has helped to determine morphological change along lines of least evolutionary resistance, influencing sexual dimorphism. In Miopithecus and Erythrocebus, the smallest and largest guenon genera, it is likely that the interaction of ecology and size contributes significantly to patterns of sexual dimorphism. The results of this study thus emphasise the need to consider allometry and size alongside ecology and behaviour when examining primate sexual dimorphism.     

Divergence in size,but not in shape: variation in skull size and shape within Ommatotriton newts

Using a geometric morphometric approach, we explored the variation in skull size and skull shape in banded newts (genus Ommatotriton). The genus Ommatotriton is represented by two allopatric, genetically well‐defined species: Ommatotriton ophryticus and O. vittatus. Within each species, two subspecies have been recognised. The samples used in this study cover the geographical and genetic variation within each species. We found statistically significant variation in skull size between species and among populations within species. When corrected for size, there was no significant variation in shape between species. Our results indicate that the variation in skull shape within the genus Ommatotriton is almost entirely due to size‐dependent, allometric shape changes. The exception is the shape of the ventral skull in males. Males of O. ophryticus and O. vittatus significantly diverge in the shape of the ventral cranium. The ventral skull, more precisely the upper jaw and palate, is directly functionally related to feeding. In general, our results indicate that allometry is a significant factor in the morphological variation of banded newts. However, the divergence in the ventral skull shape of males indicates that sexual selection and niche partitioning may have influenced the evolution of skull shape in these newts.     

Microevolutionary patterns in the common caiman predict macroevolutionary trends across extant crocodilians

Both extinct and extant crocodilians have repeatedly diversified in skull shape along a continuum, from narrow‐snouted to broad‐snouted phenotypes. These patterns occur with striking regularity, although it is currently unknown whether these trends also apply to microevolutionary divergence during population differentiation or the early stages of speciation. Assessing patterns of intraspecific variation within a single taxon can potentially provide insight into the processes of macroevolutionary differentiation. For example, high levels of intraspecific variation along a narrow‐broad axis would be consistent with the view that cranial shapes can show predictable patterns of differentiation on relatively short timescales, and potentially scale up to explain broader macroevolutionary patterns. In the present study, we use geometric morphometric methods to characterize intraspecific cranial shape variation among groups within a single, widely distributed clade, Caiman crocodilus. We show that C. crocodilus skulls vary along a narrow/broad‐snouted continuum, with different subspecies strongly clustered at distinct ends of the continuum. We quantitatively compare these microevolutionary trends with patterns of diversity at macroevolutionary scales (among all extant crocodilians). We find that morphological differences among the subspecies of C. crocodilus parallel the patterns of morphological differentiation across extant crocodilians, with the primary axes of morphological diversity being highly correlated across the two scales. We find intraspecific cranial shape variation within C. crocodilus to span variation characterized by more than half of living species. We show the main axis of intraspecific phenotypic variation to align with the principal direction of macroevolutionary diversification in crocodilian cranial shape, suggesting that mechanisms of microevolutionary divergence within species may also explain broader patterns of diversification at higher taxonomic levels.     

Sexual dimorphism in the skull geometry of newt species of <Emphasis Type="Italic">Ichthyosaura,Triturus</Emphasis> and <Emphasis Type="Italic">Lissotriton</Emphasis> (Salamandridae,Caudata, Amphibia)

In this study, we applied geometric morphometrics to explore variations in the level and pattern of sexual size dimorphism (SSD) and sexual shape dimorphism (SShD) of the ventral cranium in three different Modern Eurasian newt taxa (Ichthyosaura alpestris, Triturus species group and Lissotriton vulgaris). The ventral cranium is the part of the skull that is more directly related to foraging and feeding. Our results indicate that the level and pattern of sexual dimorphism in the ventral cranium differ among Modern Eurasian newt taxa. Regarding sexual dimorphism in skull size, Ichthyosaura alpestris and Triturus species show female-biased patterns (females are larger than males), whereas Lissotriton vulgaris appears to be non-dimorphic in skull size. In I. alpestris and Triturus species, SShD is mostly absent, whereas in L. vulgaris, SShD is more pronounced. A high level of variation between populations in both SSD and SShD indicates that local conditions may have a profound effect on the magnitude and direction of sexual dimorphism. The significant sexual differences in ventral cranium size and shape indicate possible subtle intersexual differences in ecological demands due to diet specialisation, in spite of similar general ecological settings.     

Sexual dimorphism in the canines and skulls of carnivores: effects of size, phylogency, and behavioural ecology

Sexual dimorphism in craniodental features is investigated in a sample of 45 carnivore species in relation to allometry, phylogeny, and behavioural ecology. Dimorphism is more pronounced in both upper and lower canine size and strength than in carnassial size, skull dimensions and biomechanical features, but all dimorphism indices covary. As with most morphological characters, differences in canine sexual dimorphism are significantly related to phylogeny, estimated from either taxonomic rankings or a limited matrix of molecular distances; in particular, mustelids, felids and procyonids are more dimorphic than other carnivore families. Thus, because of problems related to species dependence in comparative data, remaining analyses are based on phylogenetically transformed values using a spatial autoregressive method.
In contrast to other mammals, sexual dimorphism in carnivore canines is not correlated with differences in body weight, skull length or basicranial axis length. Nor is it correlated with categorical variables of activity pattern, habitat, or diet. In our Carnivore sample, canine dimorphism is related only to breeding system: uni-male, group-living (harem) species have significantly greater canine dimorphism than multi-male, multi-female groups and monogamous pair-bonding species. By contrast, dimorphism in carnassial size is related to dietary differences, specifically greater dimorphism in meat-eating species, and not breeding patterns. Dimorphism in estimates of jaw muscle size suggest functional demands from both diet and breeding type. It is concluded that, befitting patterns of heterodont dentition, sexual selection influences variation in canine dimorphism while feeding ecology is related to carnassial dimorphism.     

Males Resemble Females: Re-Evaluating Sexual Dimorphism in Protoceratops andrewsi (Neoceratopsia,Protoceratopsidae)

BackgroundProtoceratops andrewsi (Neoceratopsia, Protoceratopsidae) is a well-known dinosaur from the Upper Cretaceous of Mongolia. Some previous workers hypothesized sexual dimorphism in the cranial shape of this taxon, using qualitative and quantitative observations. In particular, width and height of the frill as well as the development of a nasal horn have been hypothesized as potentially sexually dimorphic.Conclusions/SignificanceSexual dimorphism within Protoceratops andrewsi is not strongly supported by our results, as previously proposed by several authors. Anatomical traits such as height and width of the frill, and skull size thus may not be sexually dimorphic. Based on PCA for a data set focusing on the rostrum and associated ANOVA results, nasal horn height is the only feature with potential dimorphism. As a whole, most purported dimorphic variation is probably primarily the result of ontogenetic cranial shape changes as well as intraspecific cranial variation independent of sex.     

Ontogenetic allometry constrains cranial shape of the head‐first burrowing worm lizard Cynisca leucura (Squamata: Amphisbaenidae)

Amphisbaenians are fossorial, predominantly limbless squamate reptiles with distinct cranial shapes corresponding to specific burrowing behaviors. Due to their cryptic lifestyles and the scarcity of museum specimens, little is known of their intraspecific variation, particularly regarding cranial osteology. This represents a critical lack of information, because the majority of morphological investigations of squamate relationships are based on cranial characters. We investigated cranial variation in the West African Coast Worm Lizard Cynisca leucura, a round‐headed member of the Amphisbaenidae. Using geometric morphometric analyses of three‐dimensional computed tomographic scans, we found that cranial osteology of C. leucura is highly conserved, with the majority of shape changes occurring during growth as the cranium becomes more slender and elongate, accompanied by increasing interdigitation among the dermal roofing bones. Elements of the ventral portion of the cranium remain loosely connected in adults, possibly as a protective mechanism against repeated compression and torsion during burrow excavation. Intraspecific variation was strongly correlated with size change from juveniles to adults, indicating a dominant role of ontogenetic allometry in determining cranial shape. We found no evidence of sexual dimorphism, either during growth or among adults. Given the fossorial habits of C. leucura, we hypothesize that cranial allometry is under strong stabilizing selection to maintain adequate proportions for head‐first digging, thereby constraining the ability of individuals to respond to differing selection pressures, including sexual selection and variation in diet or microhabitat. For species in which digging imposes less mechanical stress (e.g., in softer sand), allometric associations during growth may be weakened, allowing changes to the ontogenetic trajectory and subsequent morphological traits. Such developmental dissociation between size and shape, known as heterochrony, may also be implicit in the evolution of the other amphisbaenian cranial shapes (shovel, spade, and keel), which may themselves be functionally adapted for their respective burrowing techniques. J. Morphol. 277:1159–1167, 2016. © 2016 Wiley Periodicals, Inc.     

A complete survey of normal pores on a smooth shell ostracod (Crustacea): Landmark‐based versus outline geometric morphometrics

Pores and sensilla on ostracod shell have often been used in studies of ontogeny, taxonomy, and phylogeny of the group. However, an analysis of sexual dimorphism and variation between valves in the number and distribution of pores is lacking. Also, such studies have never been done on a widely distributed, morphologically variable, and weakly ornamented freshwater ostracod. Here, we survey pores in one such species, Physocypria kraepelini . We choose 27 homologous pores as landmarks for 2D‐geometric morphometric analysis, with the aim to assess intersexual and between valves variation in size and shape relative to the Fourier outline analysis. This species has only simple (Type A) pores with and without a lip, and each pore carries an undivided sensory seta. Our results show that the total number of pores varies (from 270 to 296), but this is not associated with a specific valve. Males carry fewer pores than females, however no sex specific pores are found. Small intrapopulation divergence of the Cyt b molecular marker (1%) indicates that morphological variability is not species related. We found that P. kraepelini exhibits directional asymmetry of size and shape, sexual size dimorphism (SSD) but lacks sexual shape dimorphism (SShD). Two geometric morphometrics methods were congruent in the estimation of SSD, SShD, and directional asymmetry of shape but differ in the statistical evaluation of directional asymmetry of size. Contrary to other animal groups, our study suggests that ostracods have more pronounced directional asymmetry of shape compared to directional asymmetry of size.     

Pollinator size and its consequences: Robust estimates of body size in pollinating insects

Body size is an integral functional trait that underlies pollination‐related ecological processes, yet it is often impractical to measure directly. Allometric scaling laws have been used to overcome this problem. However, most existing models rely upon small sample sizes, geographically restricted sampling and have limited applicability for non‐bee taxa. Allometric models that consider biogeography, phylogenetic relatedness, and intraspecific variation are urgently required to ensure greater accuracy. We measured body size as dry weight and intertegular distance (ITD) of 391 bee species (4,035 specimens) and 103 hoverfly species (399 specimens) across four biogeographic regions: Australia, Europe, North America, and South America. We updated existing models within a Bayesian mixed‐model framework to test the power of ITD to predict interspecific variation in pollinator dry weight in interaction with different co‐variates: phylogeny or taxonomy, sexual dimorphism, and biogeographic region. In addition, we used ordinary least squares regression to assess intraspecific dry weight ~ ITD relationships for ten bees and five hoverfly species. Including co‐variates led to more robust interspecific body size predictions for both bees and hoverflies relative to models with the ITD alone. In contrast, at the intraspecific level, our results demonstrate that the ITD is an inconsistent predictor of body size for bees and hoverflies. The use of allometric scaling laws to estimate body size is more suitable for interspecific comparative analyses than assessing intraspecific variation. Collectively, these models form the basis of the dynamic R package, “pollimetry,” which provides a comprehensive resource for allometric pollination research worldwide.     

Sexual dimorphism in striped dolphin (Stenella coeruleoalba) crania from South Africa

This study assessed the extent of sexual dimorphism in striped dolphin (Stenella coeruleoalba) cranial size and shape off the South African coast. Dorsal and ventral features of 60 striped dolphin crania from both the western and eastern coasts of South Africa were analyzed using landmark‐based geometric morphometrics. Although there was no evidence of dimorphism in cranial size, evidence for small, but significant, variation in both dorsal and ventral cranial shape was found between the sexes. The observed dimorphism was partly associated with changes in shape around the temporal fossa, occipital condyle and supraoccipital bone, the nasal bone, and the paraoccipital process and basiocciptal. The temporal fossa serves as an attachment point for the temporal muscle, which functions to close the lower jaw, and the occipital area serves as the anterior insertion of the epaxial muscles, which power the upstroke of the flukes during swimming. Both the paraoccipital process and basiocciptal are associated with the functioning of the hyoid apparatus, which serves as an attachment point for muscles and ligaments involved in feeding and sound production. These findings suggest the possibility of differences in diet, foraging behavior, vocalizations, and locomotion between the sexes of this species.     

Ontogenetic approaches to sexual dimorphism in anthropoids

Studies of sexual dimorphism have traditionally focused on the static differences in size and shape between adult males and females. In this paper, I suggest that an investigation of the ontogenetic bases of sexual dimorphism can provide new insights and information unobtainable from studies concerned only with adult endpoints. While growth is often viewed as simply the developmental pathway utilized to attain final adult size and shape, we must recognize that it is the entire pattern of sex-differentiated growth, and not merely the adult endpoints, which is adaptive and the target of natural selection. The importance of an ontogenetic approach to the analysis of sexual dimorphism is also demonstrated by the fact that a given morphologicalresult (e.g., a certain degree of adult weight dimorphism) may be attained by very different developmentalprocesses, signalling selection for quite different factors. The need to analyze the ontogenetic bases of sexual dimorphism in size and shape has recently been recognized by Jarman, in his study of dimorphism in large terrestrial herbivores. Here I combine aspects of Jarman’s approach with those of allometry and heterochrony in an analysis of sexual dimorphism in selected anthropoid primates. It is demonstrated that although all dimorphic anthropoids appear to be characterized by somebimaturism, the degree varies significantly. Marked weight dimorphism in certain species is primarily produced by an increased differentiation of female and male growthrates, while in other species the primary change involves differences in thetime or duration of growth between the sexes. These variations are illustrated with anthropoid genera such asMiopithecus, Cercopithecus, Erythrocebus, Macaca, Papio, Pan, andGorilla. It is suggested that additional ontogenetic investigations of other anthropoids will help clarify some of the socioecological bases of this variation in the ways of attaining an adult dimorphic state. This will contribute to our understanding of the complex factors underlying and producing sexual dimorphism in primates and other mammals.     

Cranial shape variation in jacarean caimanines (Crocodylia,Alligatoroidea) and its implications in the taxonomic status of extinct species: The case of Melanosuchus fisheri

Melanosuchus niger (Crocodylia, Alligatoroidea) is one of the six living caimanine species widely distributed throughout the Amazon River basin today. Although there is only one extant species of Melanosuchus, fossil material assigned to this genus, represented by M. fisheri, has been reported from the late Miocene in South America. However, the validity of this taxon has been questioned and a recent investigation indicates that the referred specimen of M. fisheri (MCZ 4336) actually belongs to Globidentosuchus brachyrostris, while those diagnostic characters present in the holotype (MCNC 243) fall into the spectrum of intraspecific variation of M. niger. Here, we compare the skull shape of the holotype of M. fisheri with the ontogenetic series of the four jacarean species (M. niger, Caiman yacare, Caiman crocodilus, and Caiman latirostris) using 2D‐geometric morphometric analyses in two different views. The analyses indicate that MCNC 243 falls into the morphospace of M. niger and C. latirostris. Despite strong shape similarities between juveniles of C. latirostris and MCNC 243, further anatomical comparisons reveal notable differences between them. In contrast, no concrete anatomical differences can be found between MCNC 243 and M. niger, although shape analyses indicate that MCNC 243 is relatively robust for its size. Thus, this study is able to confirm that the genus Melanosuchus was present in the late Miocene, but it still remains unclear if MCNC 243 should be treated as a junior synonym or probably a sister species of M. niger. Its Miocene age favors the second option, but as the shape analyses were also not able to extract any diagnostic characters, it should be retained as Melanosuchus cf. niger.     

Ontogenetic allometry, heterochrony, and interspecific differences in the skull of African apes, using tridimensional Procrustes analysis

Ontogenetic studies of African ape skulls lead to an analysis of morphological differences in terms of allometry, heterochrony, and sexual dimorphism. The use of geometric morphometrics allows us 1) to define size and shape variations as independent factors (an essential but seldom respected condition for heterochrony), and 2) to calculate in percentage of shape changes and to graphically represent the parts of shape variation which are related to various biological phenomena: common allometry, intraspecific allometry, and allometric and nonallometric shape discrimination. Three tridimensional Procrustes analyses and the calculation of multivariate allometries, discriminant functions, and statistical tests are used to compare the skulls of 50 Pan troglodytes, and 50 Gorilla gorilla of different dental stages. The results both complement and modify classical results obtained from similar material but with different methods. Size and Scaling in Primate Morphology, New York: Plenum, p. 175-205). As previously described by Shea, the common growth allometric pattern is very important (64% of total shape variation). It corresponds to a larger increase of facial volume than of neurocranial volume, a more obliquely oriented foramen magnum, and a noticeable reshaping of the nuchal region (higher inion). However, the heterochronic interpretation based on common allometry is rather different from Shea. Gorillas differ from chimpanzees not only with a larger magnitude of allometric change (rate peramorphosis), as is classically said, but also grow more in size than in shape (size acceleration). In other words, for a similar stage of growth, gorillas have the size and shape corresponding to older chimpanzees, and for a similar shape, gorillas have a larger size than chimpanzees. In contrast, sexual dimorphism actually corresponds to allometric changes only, as classically demonstrated (time hypermorphosis). Sexual dimorphism is here significant in adult gorillas alone, and solely in terms of allometry (size-related shape and size, given that sagittal and nuchal crests are not taken into account). The study also permits us to differentiate two different shape variations that are classically confused in ontogenetic studies: a very small part of allometric shape change which is specific to each species (1% of the total shape variation), and nonallometric species-specific traits independent of growth (8% of total shape change). When calculated in terms of intraspecific allometries (including common allometry and noncommon allometry), shape changes are more extensive in gorillas (36% of total shape change) than in chimpanzees (29% of total shape change). The allometric differences mainly concern the inion, which becomes higher; the position of the foramen magnum, more dorsally oriented; and the palate, more tilted in adult gorillas than in adult chimpanzees. In contrast, nonallometric species-specific traits in gorillas are the long and flat vault characterized by a prominent occipital region, the higher and displaced backward glabella, and the protrusive nose. Biomechanical schemes built from shape partition suggest that the increased out-of-plumb position of the head during growth is partially compensated in gorillas by a powerful nuchal musculature due to the peculiar shape of the occipital region.