Relationships among ontogenetic,static, and evolutionary allometry

The relationship between ontogenetic, static, and evolutionary levels of allometry is investigated. Extrapolation from relative size relationships in adults to relative growth in ontogeny depends on the variability of slopes and intercepts of ontogenetic vectors relative to variability in length of the vector. If variability in slopes and intercepts is low relative to variability in length, ontogenetic and static allometries will be similar. The similarity of ontogenetic and static allometries was tested by comparing the first principal component, or size vector, for correlations among 48 cranial traits in a cross-sectional ontogenetic sample of rhesus macaques from Cayo Santiago with a static sample from which all age- and sex-related variation had been removed. The vector correlation between the components is high but significantly less than one while two of three allometric patterns apparent in the ontogenetic component are not discernable in the static component. This indicates that there are important differences in size and shape relationships among adults and within ontogenies. Extrapolation from intra- or interspecific phenotypic allometry to evolutionary allometry is shown to depend on the similarity of genetic and phenotypic allometry patterns. Similarity of patterns was tested by comparing the first principal components of the phenotypic, genetic, and environmental correlation matrices calculated using standard quantitative genetic methods. The patterns of phenotypic, genetic, and environmental allometry are dissimilar; only the environmental allometries show ontogenetic allometric patterns. This indicates that phenotypic allometry may not be an accurate guide to patterns of evolutionary change in size and shape.     

EVOLUTION OF STATIC ALLOMETRIES: ADAPTIVE CHANGE IN ALLOMETRIC SLOPES OF EYE SPAN IN STALK‐EYED FLIES

Julian Huxley showed that within‐species (static) allometric (power‐law) relations can arise from proportional growth regulation with the exponent in the power law equaling the factor of proportionality. Allometric exponents may therefore be hard to change and act as constraints on the independent evolution of traits. In apparent contradiction to this, many empirical studies have concluded that static allometries are evolvable. Many of these studies have been based, however, on a broad definition of allometry that includes any monotonic shape change with size, and do not falsify the hypothesis of constrained narrow‐sense allometry. Here, we present the first phylogenetic comparative study of narrow‐sense allometric exponents based on a reanalysis of data on eye span and body size in stalk‐eyed flies (Diopsidae). Consistent with a role in sexual selection, we found strong evidence that male slopes were tracking “optima” based on sexual dimorphism and relative male trait size. This tracking was slow, however, with estimated times of 2–3 million years for adaptation to exceed ancestral influence on the trait. Our results are therefore consistent with adaptive evolution on million‐year time scales, but cannot rule out that static allometry may act as a constraint on eye‐span adaptation at shorter time scales.     

Evolution of extreme ontogenetic allometric diversity and heterochrony in pythons,a clade of giant and dwarf snakes

Ontogenetic allometry, how species change with size through their lives, and heterochony, a decoupling between shape, size, and age, are major contributors to biological diversity. However, macroevolutionary allometric and heterochronic trends remain poorly understood because previous studies have focused on small groups of closely related species. Here, we focus on testing hypotheses about the evolution of allometry and how allometry and heterochrony drive morphological diversification at the level of an entire species‐rich and diverse clade. Pythons are a useful system due to their remarkably diverse and well‐adapted phenotypes and extreme size disparity. We collected detailed phenotype data on 40 of the 44 species of python from 1191 specimens. We used a suite of analyses to test for shifts in allometric trajectories that modify morphological diversity. Heterochrony is the main driver of initial divergence within python clades, and shifts in the slopes of allometric trajectories make exploration of novel phenotypes possible later in divergence history. We found that allometric coefficients are highly evolvable and there is an association between ontogenetic allometry and ecology, suggesting that allometry is both labile and adaptive rather than a constraint on possible phenotypes.     

The ontogeny of sexual dimorphism in the cranium of Bornean orang-utans (Pongo pygmaeus pygmaeus): II. Allometry and heterochrony

Based on a homogeneous sample of 212 individuals spanning all postnatal periods, we examine the ontogeny of cranial sexual dimorphism in Bornean orang-utans (Pongo pygmaeus pygmaeus) by means of allometric analysis and in terms of heterochrony. The bivariate growth allometries of 20 cranial dimensions against basicranial length yield two major patterns. Confirming the null hypothesis, strong ontogenetic scaling, where growth regressions of both sexes fall along a single ontogenetic continuum, and where shape differences between adult males and females result from the extension of relative growth in the smaller females to larger size in males, is found in 10 cases. Ontogenetic scaling is particularly strong in proportions of (1) the neurocranium directly associated with brain size, (2) the orbital region, and (3) the dental arcade. In terms of heterochrony such a pattern most likely is the result of a process termed "time hypermorphosis", i.e. an extension of the growth period in time in males. The second major pattern seen in the remaining 10 cases shows a departure from ontogenetic scaling, with males exhibiting a significantly steeper slope than females. Departures from ontogenetic scaling, where size and shape are dissociated with adult males being disproportionately larger than adult females, are found in proportions of cranial regions directly associated with secondary sexual character development: prognathism, canine size, and cheek pad area. In terms of heterochrony such a pattern most likely is the result of a process termed "acceleration", i.e. the rate of shape change is increased in males.     

草兔胚后发育过程中头骨大小与形状变化的几何形态学分析(兔形目:兔科)

在本文中,我们使用几何形态学的研究方法对58 个处于不同年龄阶段的草兔头骨进行了分析。分析中总共使用来自头骨背面、腹面和侧面的180 个标点和半标点。研究结果表明,草兔在胚后发育早期即迅速建立起与成年个体近似的形态结构,组成头骨的不同形态单元存在显著的异速生长现象,主要的形变发生在幼年至年龄1 阶段,即出生后的6 个月以内。从大小的变化来看,鼻骨在胚后发育过程中呈现正的异速生长,额骨和眼眶区则与头骨整体大小变化基本等速,而顶骨、听泡和枕骨大孔则呈现显著的负异速生长。我们亦用几何形态学的方法绘制出头骨在生长发育早期和晚期不同部位的形态变化轨迹,这一结果显示较显著的形状变化发生在鼻骨、前颌骨、眶上突和头骨的纵轴方向。头骨整体形态在胚后发育过程中伸长并变窄。这些变化将有利于幼兔较早实现完善的头部系统的建立,尤其有利于提高呼吸系统的通风能力,提高在高强度的捕食压力下保持警觉,在多样的运动过程中保持身体平稳的能力,也可能进一步提高了对固态食物的处理能力。这种异速生长模式可能是善于奔跑的植食性哺乳动物在功能需求上的一种适应性进化特征。     

Allometry and heterochrony in the African apes

In this work allometry and heterochrony are integrated in an analysis of ontogenic and interspecific morphological patterns in the African apes. The relationship between the interspecific differences in adult morphology and the differences in underlying patterns of growth allometries, body weight growth rates, and developmental chronologies is investigated. Results indicate that rate hypermorphosis, or the extension of ancestral allometries into new size/shape ranges with no increase in the duration of ontogeny, underlies many of the interspecific differences in form among the African apes. In addition, the need for further clarification of the processes of heterochrony is stressed by distinguishing between rate and timing differences. These distinctions and processes are illustrated and discussed using the morphological data on the African apes.     

ALLOMETRIC CONSTRAINTS AND THE EVOLUTION OF ALLOMETRY

Morphological traits often covary within and among species according to simple power laws referred to as allometry. Such allometric relationships may result from common growth regulation, and this has given rise to the hypothesis that allometric exponents may have low evolvability and constrain trait evolution. We formalize hypotheses for how allometry may constrain morphological trait evolution across taxa, and test these using more than 300 empirical estimates of static (within‐species) allometric relations of animal morphological traits. Although we find evidence for evolutionary changes in allometric parameters on million‐year, cross‐species time scales, there is limited evidence for microevolutionary changes in allometric slopes. Accordingly, we find that static allometries often predict evolutionary allometries on the subspecies level, but less so across species. Although there is a large body of work on allometry in a broad sense that includes all kinds of morphological trait–size relationships, we found relatively little information about the evolution of allometry in the narrow sense of a power relationship. Despite the many claims of microevolutionary changes of static allometries in the literature, hardly any of these apply to narrow‐sense allometry, and we argue that the hypothesis of strongly constrained static allometric slopes remains viable.     

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.     

Assessing endocranial variations in great apes and humans using 3D data from virtual endocasts

Modern humans are characterized by their large, complex, and specialized brain. Human brain evolution can be addressed through direct evidence provided by fossil hominid endocasts (i.e. paleoneurology), or through indirect evidence of extant species comparative neurology. Here we use the second approach, providing an extant comparative framework for hominid paleoneurological studies. We explore endocranial size and shape differences among great apes and humans, as well as between sexes. We virtually extracted 72 endocasts, sampling all extant great ape species and modern humans, and digitized 37 landmarks on each for 3D generalized Procrustes analysis. All species can be differentiated by their endocranial shape. Among great apes, endocranial shapes vary from short (orangutans) to long (gorillas), perhaps in relation to different facial orientations. Endocranial shape differences among African apes are partly allometric. Major endocranial traits distinguishing humans from great apes are endocranial globularity, reflecting neurological reorganization, and features linked to structural responses to posture and bipedal locomotion. Human endocasts are also characterized by posterior location of foramina rotunda relative to optic canals, which could be correlated to lesser subnasal prognathism compared to living great apes. Species with larger brains (gorillas and humans) display greater sexual dimorphism in endocranial size, while sexual dimorphism in endocranial shape is restricted to gorillas, differences between males and females being at least partly due to allometry. Our study of endocranial variations in extant great apes and humans provides a new comparative dataset for studies of fossil hominid endocasts.     

Developmental constraints on the evolution of wing-body allometry in Manduca sexta

Artificial selection on body size in Manduca sexta produced genetic strains with large and small body sizes. The wing-body allometries of these strains differed significantly from the wild type. Selection on small body size led to a change in the scaling of wing and body size without changing the allometry: the wings were smaller relative to the body, but to the same degree at all body sizes. Selection for large body size led to a change in allometry with a decrease in the allometric coefficient, wing size becoming progressively smaller relative to body as body size increased. When larvae were deprived of food so as to produce adults of a range of small body sizes, all strains retained the same allometric coefficient but showed an increase in the scaling factor. Thus individuals starved as larvae had a smaller adult body size but had proportionally larger wings than fed individuals. We analyzed the developmental processes that could give rise to this pattern of allometries. Differences in the relative growth of body and wing disks can account for the differences in the allometric coefficients among the three body size strains. The change in wing-body allometry at large body sizes was primarily due to an insufficient time period for growth. The available time period for growth of the wing imaginal disks poses a significant constraint on the proportional growth of wings, and thus on the evolution of large body size.     

Ontogenetic study of the skull in modern humans and the common chimpanzees: neotenic hypothesis reconsidered with a tridimensional Procrustes analysis

Heterochronic studies compare ontogenetic trajectories of an organ in different species: here, the skulls of common chimpanzees and modern humans. A growth trajectory requires three parameters: size, shape, and ontogenetic age. One of the great advantages of the Procrustes method is the precise definition of size and shape for whole organs such as the skull. The estimated ontogenetic age (dental stages) is added to the plot to give a graphical representation to compare growth trajectories. We used the skulls of 41 Homo sapiens and 50 Pan troglodytes at various stages of growth. The Procrustes superimposition of all specimens was completed by statistical procedures (principal component analysis, multivariate regression, and discriminant function) to calculate separately size-related shape changes (allometry common to chimpanzees and humans), and interspecific shape differences (discriminant function). The results confirm the neotenic theory of the human skull (sensu Gould [1977] Ontogeny and Phylogeny, Cambridge: Harvard University Press; Alberch et al. [1979] Paleobiology 5:296-317), but modify it slightly. Human growth is clearly retarded in terms of both the magnitude of changes (size-shape covariation) and shape alone (size-shape dissociation) with respect to the chimpanzees. At the end of growth, the adult skull in humans reaches an allometric shape (size-related shape) which is equivalent to that of juvenile chimpanzees with no permanent teeth, and a size which is equivalent to that of adult chimpanzees. Our results show that human neoteny involves not only shape retardation (paedomorphosis), but also changes in relative growth velocity. Before the eruption of the first molar, human growth is accelerated, and then strongly decelerated, relative to the growth of the chimpanzee as a reference. This entails a complex process, which explains why these species reach the same overall (i.e., brain + face) size in adult stage. The neotenic traits seem to concern primarily the function of encephalization, but less so other parts of the skull. Our results, based on the discriminant function, reveal that additional structural traits (corresponding to the nonallometric part of the shape which is specific to humans) are rather situated in the other part of the skull. They mainly concern the equilibrium of the head related to bipedalism, and the respiratory and masticatory functions. Thus, the reduced prognathism, the flexed cranial base (forward position of the foramen magnum which is brought closer to the palate), the reduced anterior portion of the face, the reduced glabella, and the prominent nose mainly correspond to functional innovations which have nothing to do with a neotenic process in human evolution. The statistical analysis used here gives us the possibility to point out that some traits, which have been classically described as paedomorphic because they superficially resemble juvenile traits, are in reality independent of growth.     

Sexual selection and allometry: a critical reappraisal of the evidence and ideas

One of the most pervasive ideas in the sexual selection literature is the belief that sexually selected traits almost universally exhibit positive static allometries (i.e., within a sample of conspecific adults, larger individuals have disproportionally larger traits). In this review, I show that this idea is contradicted by empirical evidence and theory. Although positive allometry is a typical attribute of some sexual traits in certain groups, the preponderance of positively allometric sexual traits in the empirical literature apparently results from a sampling bias reflecting a fascination with unusually exaggerated (bizarre) traits. I review empirical examples from a broad range of taxa illustrating the diversity of allometric patterns exhibited by signal, weapon, clasping and genital traits, as well as nonsexual traits. This evidence suggests that positive allometry may be the exception rather than the rule in sexual traits, that directional sexual selection does not necessarily lead to the evolution of positive allometry and, conversely, that positive allometry is not necessarily a consequence of sexual selection, and that many sexual traits exhibit sex differences in allometric intercept rather than slope. Such diversity in the allometries of secondary sexual traits is to be expected, given that optimal allometry should reflect resource allocation trade-offs, and patterns of sexual and viability selection on both trait size and body size. An unbiased empirical assessment of the relation between sexual selection and allometry is an essential step towards an understanding of this diversity.     

Does allometry account for shape variability in <Emphasis Type="Italic">Ephedrus persicae</Emphasis> Froggatt (Hymenoptera: Braconidae: Aphidiinae) parasitic wasps?

We analysed linear measurements on various parts of the body and the configuration of 11 landmarks on the wing in a large sample of Ephedrus persicae that had emerged from 13 aphid host species, to assess whether static allometry (a measure of the scaling relationship between traits in a population of individuals at the same ontogenetic stage) accounts for variation in body shape. The analysed specimens came from several localities in Europe, Asia Minor, Japan and South America, and cover a large portion of the distribution area of E. persicae. We found that allometry accounts for variation in body shape among different biotypes within the E. persicae group. The allometric slopes for head size (HD), petiolus width (PETW), mesoscutum width (MSC), and ovipositor sheath length (OVPL) diverged significantly among biotypes, indicating biotype-specific allometries. The analysis of allometric variation in wing shape showed that the pattern and direction of allometric changes also differed among individuals that had emerged from different hosts. Our results (observed divergences in the directions of allometric slopes of particular morphometric traits and wing shape) suggest that allometric relations within E. persicae are not conserved, so that allometry itself changes, evolving differently in aphid parasitoids that emerge from different hosts.     

Allometric and nonallometric components of Drosophila wing shape respond differently to developmental temperature

Phenotypic plasticity of wing size and shape of Drosophila simulans was analyzed across the entire range of viable developmental temperatures with Procrustes geometric morphometric method. In agreement with previous studies, size clearly decreases when temperature increases. Wing shape variation was decomposed into its allometric (24%) and nonallometric (76%) components, and both were shown to involve landmarks located throughout the entire wing blade. The allometric component basically revealed a progressive, monotonous variation along the temperature. Surprisingly, nonallometric shape changes were highly similar at both extremes of the thermal range, suggesting that stress, rather than temperature per se, is the key developmental factor affecting wing shape.     

Crown allometries are less responsive than stem allometry to tree size and habitat variations in an Indian monsoon forest

While theoretical allometric models postulate universal scaling exponents, empirical relationships between tree dimensions show marked variability that reflects changes in the biomass allocation pattern. As growth of the various tree compartments may be controlled by different functions, it is hypothesized that they may respond differently to factors of variation, resulting in variable tree morphologies and potentially in trade-offs between allometric relationships. We explore the variability of tree stem and crown allometries using a dataset of 1,729 trees located in an undisturbed wet evergreen forest of the Western Ghats, India. We specifically test whether species adult stature, terrain slope, tree size and crown light exposure affect the relationships between stem diameter and stem height (stem allometry), and between stem diameter and crown width, crown area and crown volume (crown allometries). Results show that both stem and crown allometries are subject to variations in relation to both endogenous (tree size, species adult stature) and exogenous (terrain slope, crown light exposure) factors. Stem allometry appears to be more affected by these factors than are crown allometries, including the stem diameter–crown volume relationship, which proved to be particularly stable. Our results support the idea that height is a prevailing adjustment factor for a tree facing variable growth (notably light) conditions, while stem diameter–crown volume allometry responds more to internal metabolic constraints. We ultimately discuss the various sources of variability in the stem and crown allometries of tropical trees that likely play an important role in forest community dynamics.     

Variation in the ontogenetic allometry of horn length in bovids along a body mass continuum

Allometric relationships describe the proportional covariation between morphological, physiological, or life‐history traits and the size of the organisms. Evolutionary allometries estimated among species are expected to result from species differences in ontogenetic allometry, but it remains uncertain whether ontogenetic allometric parameters and particularly the ontogenetic slope can evolve. In bovids, the nonlinear evolutionary allometry between horn length and body mass in males suggests systematic changes in ontogenetic allometry with increasing species body mass. To test this hypothesis, we estimated ontogenetic allometry between horn length and body mass in males and females of 19 bovid species ranging from ca. 5 to 700 kg. Ontogenetic allometry changed systematically with species body mass from steep ontogenetic allometries over a short period of horn growth in small species to shallow allometry with the growth period of horns matching the period of body mass increase in the largest species. Intermediate species displayed steep allometry over long period of horn growth. Females tended to display shallower ontogenetic allometry with longer horn growth compared to males, but these differences were weak and highly variable. These findings show that ontogenetic allometric slope evolved across species possibly as a response to size‐related changes in the selection pressures acting on horn length and body mass.     

Heterochrony and allometry: the analysis of evolutionary change in ontogeny

The connection between development and evolution has become the focus of an increasing amount of research in recent years, and heterochrony has long been a key concept in this relation. Heterochrony is defined as evolutionary change in rates and timing of developmental processes; the dimension of time is therefore an essential part in studies of heterochrony. Over the past two decades, evolutionary biologists have used several methodological frameworks to analyse heterochrony, which differ substantially in the way they characterize evolutionary changes in ontogenies and in the resulting classification, although they mostly use the same terms. This review examines how these methods compare ancestral and descendant ontogenies, emphasizing their differences and the potential for contradictory results from analyses using different frameworks. One of the two principal methods uses a clock as a graphical display for comparisons of size, shape and age at a particular ontogenic stage, whereas the other characterizes a developmental process by its time of onset, rate, and time of cessation. The literature on human heterochrony provides particularly clear examples of how these differences produce apparent contradictions when applied to the same problem. Developmental biologists recently have extended the concept of heterochrony to the earliest stages of development and have applied it at the cellular and molecular scale. This extension brought considerations of developmental mechanisms and genetics into the study of heterochrony, which previously was based primarily on phenomenological characterizations of morphological change in ontogeny. Allometry is the pattern of covariation among several morphological traits or between measures of size and shape; unlike heterochrony, allometry does not deal with time explicitly. Two main approaches to the study of allometry are distinguished, which differ in the way they characterize organismal form. One approach defines shape as proportions among measurements, based on considerations of geometric similarity, whereas the other focuses on the covariation among measurements in ontogeny and evolution. Both are related conceptually and through the use of similar algebra. In addition, there are close connections between heterochrony and changes in allometric growth trajectories, although there is no one-to-one correspondence. These relationships and outline links between different analytical frameworks are discussed.     

Shape Variation in the Dermatocranium of the Greater Short-Horned Lizard <Emphasis Type="Italic">Phrynosoma hernandesi</Emphasis> (Reptilia: Squamata: Phrynosomatidae)

Dermatocranial shape and horn morphology display great disparity among the species of Phrynosoma. Ontogenetic change in dermatocranial shape in a series of 79 specimens of the short-horned Phrynosoma hernandesi (54F: 25M) was examined using geometric morphometric techniques. A multivariate ANCOVA of Procrustes residuals with sex as a factor and ln(centroid size) as the covariate indicated sexual shape dimorphism. Separate multivariate regressions of Procrustes residuals on ln(centroid size) for each sex indicated that allometry accounts for ~52–54% of the total sample shape variance. Comparisons of ontogenetic shape change between sexes indicate that sexual shape dimorphism is minimal and of uncertain biological significance. Groupings of multivariate regression coefficients by magnitude and sign suggest that allometric integration of the dermatocranium is not uniform over the dermatocranium. Principal component analysis of the landmark configurations corrected for sex and allometry yields a first principal component which describes shape variance concentrated in the posterolateral and posterior regions of the dermatocranium, and again is indicative of non-uniform shape variation over the dermatocranium. Our findings for P. hernandesi indicate that the adult shape of the dermatocranium may contribute to a passive defence against predation. We hypothesize that the complexity in dermatocranial shape demonstrated here for P. hernandesi indicates parcellation of shape variance, which may contribute to explanations of the pronounced dermatocranial disparity exhibited by the species of Phrynosoma.     

Sexual selection explains Rensch's rule of allometry for sexual size dimorphism

In 1950, Rensch first described that in groups of related species, sexual size dimorphism is more pronounced in larger species. This widespread and fundamental allometric relationship is now commonly referred to as 'Rensch's rule'. However, despite numerous recent studies, we still do not have a general explanation for this allometry. Here we report that patterns of allometry in over 5300 bird species demonstrate that Rensch's rule is driven by a correlated evolutionary change in females to directional sexual selection on males. First, in detailed multivariate analysis, the strength of sexual selection was, by far, the strongest predictor of allometry. This was found to be the case even after controlling for numerous potential confounding factors, such as overall size, degree of ornamentation, phylogenetic history and the range and degree of size dimorphism. Second, in groups where sexual selection is stronger in females, allometry consistently goes in the opposite direction to Rensch's rule. Taken together, these results provide the first clear solution to the long-standing evolutionary problem of allometry for sexual size dimorphism: sexual selection causes size dimorphism to correlate with species size.     

Scapula form and locomotion in chimpanzee evolution

A number of primatologists have followed Coolidge (Am. J. Phys. Anthropol. 18:1–57, 1933) in suggesting that 1) there are significant shape differences in scapula form between pygmy and common chimpanzees, 2) scapulae of P. paniscus resemble those of hylobatids more than do those of P. troglodytes, and 3) therefore pygmy chimpanzees may exhibit a greater component of arm-swinging and other arboreal behaviors than common chimpanzees. In this paper I utilize a comparative analysis of ontogenetic allometries of linear dimensions to determine shape differences in the scapulae of adult pygmy and common chimpanzees and to clarify size-related changes in shape resulting from ontogenetic scaling, i.e., the differential extension of common patterns of growth allometry. Results demonstrate that the scapulae of adult P. paniscus are relatively narrower (in a direction approximately perpendicular to the scapula spine) than those of P. troglodytes, supporting Coolidge's original claim. The allometric analysis further demonstrates, however, that the two chimpanzee species exhibit ontogenetic scaling for all proportions of the scapula examined. Thus, adult pygmy chimpanzees have the scapula proportions observed in small adult and subadult P. troglodytes of comparable scapula size. The implications of this finding for past claims concerning differences in locomotor behavior between the species are discussed. This work lends additional support to previous studies that have demonstrated a high frequency of ontogenetic scaling within the genus Pan and a pedomorphic or juvenilized morphology in the pygmy chimpanzee.