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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 (Shea [1983] Am. J. Phys. Anthropol. 62:275-289; Shea [1983] Folia Primatol. (Basel) 40:32-68; Shea [1985] 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. KEY WORDS Pan troglodytes; Gorilla gorilla; skull; Procrustes; allometry; heterochrony; sexual dimorphism

KEY WORDS Procrustes superimposition; Pan troglodytes; Homo sapiens; heterochrony; allometry 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.

Changes in pelvic shape in human ontogeny and hominid phylogeny suggest that the heterochronic processes involved differ greatly from the neotenic process traditionally described in the evolution of the skull. The morphology of 150 juvenile and adult pelves of African apes, 60 juvenile and adult pelves of modern humans, two adult pelves and a juvenile hip bone of australopithecines (Sts 14, AL 288, MLD 7) was studied. Multivariate results, ontogenetic allometries, and growth curves confirm that the pelvic growth pattern in humans differs markedly from those of the African apes. The results permit the following conclusions. First, the appearance of a new feature (acetabulo-cristal buttress and cristal tubercle) at the time of human birth allows the addition of traits, such as the attainment of a proportionally narrower pelvis, with more sagittally positioned iliac blades. Pelvic proportions and orientation change progressively in early childhood as bipedalism is practiced. Other changes in pelvic proportions occur later with the adolescent growth spurt. Second, comparison of juvenile and adult australopithecines to modern humans indicates that 1) some pelvic traits of adult Australopithecus resemble those of neonate Homo; 2) the pelvic growth of Australopithecus was probably closer to that of apes, than to that of humans; and 3) prolonged growth in length of hindlimb and pelvis after sexual maturity seems to be a unique feature of Homo. The position of the acetabulo-cristal buttress and of the cristal tubercle on the ilium are similar in adult Australopithecus and neonate Homo suggesting that this feature may have been displaced later during hominid evolution. Progressive displacement of the acetabulo-cristal buttress on the ilium occurs both during hominid evolution (from Australopithecus to Homo sapiens) and human growth (from neonate to adult). This suggests peramorphic evolution of the pelvic morphology of hominids combining three processes of recapitulation (pre-displacement, acceleration and time hypermorphosis). The results lend credence to the hypothesis that no single heterochronic process accounts for all human evolutionary change; rather this reflects a combination of relative changes in growth rhythm and duration, including other perturbations, such as the appearance of new morphological features. KEY WORDS pelvis; ontogeny; heterochrony; allometry; Homo; Australopithecus; Pan; Gorilla