Size and shape dimorphism in great ape mandibles and implications for fossil species recognition

Sexual dimorphism is an important source of morphological variation, and species differences in dimorphism may be reflected in magnitude, pattern, or both. While the extant great apes are commonly used as a reference sample for distinguishing between sexual dimorphism and intertaxic variation in the fossil record, few studies have evaluated mandibular dimorphism in these taxa. In this study, percentage, degree, and pattern of mandibular dimorphism are evaluated in Pongo, Gorilla, and Pan. Mandibular dimorphism patterns are explored to determine the extent to which such patterns accurately track great ape phylogeny. Pattern stability is assessed to determine whether there are stable patterns of mandibular size and shape dimorphism that may be usefully applied to hominoid or hominid fossil species recognition studies. Finally, the established patterns of dimorphism are used to address recent debates surrounding great ape taxonomy. Results demonstrate that mandibular dimorphism is universally expressed in size, but only Pongo and Gorilla exhibit shape dimorphism. Pattern similarity tends to be greater between subspecies of the same species than between higher-order taxa, suggesting that within the great apes, there is a relationship between dimorphism pattern and phylogeny. However, this relationship is not exact, given that dimorphism patterns are weakly correlated between some closely related taxa, while great ape subspecies may be highly correlated with taxa belonging to other species or genera. Furthermore, dimorphism patterns are not significantly correlated between great ape genera, even between Gorilla and Pan. Dimorphism patterns are more stable in Gorilla and Pongo as compared to Pan, but there is little pattern stability between species or genera. Importantly, few variables differ significantly between taxa that simultaneously show consistently relatively low levels of dimorphism and low levels of variation within taxa. Combined, these findings indicate that mandibular dimorphism patterns can and do vary considerably, even among closely related species, and suggest that it would be difficult to employ great ape mandibular dimorphism patterns for purposes of distinguishing between intra- and interspecies variation in fossil samples. Finally, the degree of pattern similarity in mandibular dimorphism is lower than previously observed by others for craniofacial dimorphism. Thus, the possibility cannot be ruled out that patterns of craniofacial dimorphism in great apes may be associated with a stronger phylogenetic signal than are patterns of mandibular dimorphism.     

Scaling relationships between craniofacial sexual dimorphism and body mass dimorphism in primates: implications for the fossil record

Craniofacial remains (the most abundant identifiable remains in the fossil record) potentially offer important information about body size dimorphism in extinct species. This study evaluates the scaling relationships between body mass dimorphism and different measures of craniofacial dimorphism, evaluating taxonomic differences in the magnitude and scaling of craniofacial dimorphism across higher taxonomic groups. Data on 40 dimensions from 129 primate species and subspecies demonstrate that few dimensions change proportionally with body mass dimorphism. Primates show general patterns of greater facial vs. neurocranial and orbital dimorphism, and greater dimorphism in lengths as opposed to breadths. Within any species, though, different craniofacial dimensions can yield very different reconstructions of size dimorphism. There are significant taxonomic differences in the relationships between size and craniofacial dimorphism among primate groups that can have a significant impact on reconstructions of body mass dimorphism. Hominoids tend to show lower degrees of facial dimorphism proportional to size dimorphism than other primates. This in turn implies that strong craniofacial dimorphism in Australopithecus africanus could imply very strong body size dimorphism, conflicting with the relatively modest size dimorphism inferred from postcrania. Different methods of estimating the magnitude of size dimorphism from craniofacial measurements yield similar results, and yield comparatively low percent prediction errors for a number of dimensions. However, confidence intervals for most estimates are so large as to render most estimates highly tentative.     

The Macroevolution of our Ancient Lineage: What We Know (or Think We Know) about Early Hominin Diversity

Quantitative, evolutionary models that incorporate within- and between-species variation are critical for interpreting the fossil record of human diversity, and for making taxonomic distinctions. However, small sample sizes, sexual dimorphism, temporal trends, geographic variation, and the limited number of relevant extant models have always made the consideration of variation difficult for paleoanthropologists. Here we provide a brief overview of current early hominin diversity. We then argue that for many species our limited understanding of within species variation hampers our ability to make taxonomic decisions with any level of statistical certainty. Perhaps more significantly, the underlying causes of between-species variation among early hominins are poorly studied. There have been few attempts to correlate aspects of the phenotype with meaningful evidence for niche differentiation, to demonstrate the selective advantage of traits, or to provide other evidence for macroevolutionary divergence. Moreover, current depictions of vast pattern (but not size) diversity are inconsistent with expectations derived from most other extant primate clades that have adaptively radiated. If indeed the early hominin record is highly speciose, the reasons for this remain unclear.     

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.     

Sexual dimorphism in primates: Where are we and where do we go from here?

There are currently several debates taking place in palaeoanthropological circles in which the issue of sexual dimorphism is crucial. During the 60th Jubilee of the discovery of the TaungAustralopithecus skull, the authors concurred that much of the debate was due to differences in perception concerning dimorphism, and it was suggested that a study session dealing principally with sexual dimorphism in Primates should be organised with a view to determining what, if anything, could be observed in extant primates that might be utilised for determining the existence of dimorphism in fossil species. An advertisement was placed in the Journal of Human Evolution, calling for papers. Response was encouraging, suggesting that there was indeed a need for such a meeting, which was organised to take place late in November in Italy. Seventeen scientists sent abstracts, but due to the rather short notice, several were unable to attend the meeting. Nevertheless, papers were received and read at the meeting which form the basis for this issue of the Journal of Human Evolution. What remains to be done now, is for a second meeting to be organised at which the findings of the first meeting can be extended backwards in time into the fossil record.     

Developmental processes and canine dimorphism in primate evolution

Understanding the evolutionary history of canine sexual dimorphism is important for interpreting the developmental biology, socioecology and phylogenetic position of primates. All current evidence for extant primates indicates that canine dimorphism is achieved through bimaturism rather than via differences in rates of crown formation time. Using incremental growth lines, we charted the ontogeny of canine formation within species of Eocene Cantius, the earliest known canine-dimorphic primate, to test whether canine dimorphism via bimaturism was developmentally canalized early in primate evolution. Our results show that canine dimorphism in Cantius is achieved primarily through different rates of crown formation in males and females, not bimaturism. This is the first demonstration of rate differences resulting in canine dimorphism in any primate and therefore suggests that canine dimorphism is not developmentally homologous across Primates. The most likely interpretation is that canine dimorphism has been selected for at least twice during the course of primate evolution. The power of this approach is its ability to identify underlying developmental processes behind patterns of morphological similarity, even in long-extinct primate species.     

The evolution of body size in extant groups of North American freshwater fishes: speciation,size distributions,and Cope's rule

Change in body size within an evolutionary lineage over time has been under investigation since the synthesis of Cope's rule, which suggested that there is a tendency for mammals to evolve larger body size. Data from the fossil record have subsequently been examined for several other taxonomic groups to determine whether they also displayed an evolutionary increase in body size. However, we are not aware of any species-level study that has investigated the evolution of body size within an extant continental group. Data acquired from the fossil record and data derived from the evolutionary relationships of extant species are not similar, with each set exhibiting both strengths and weaknesses related to inferring evolutionary patterns. Consequently, expectation that general trends exhibited in the fossil record will correspond to patterns in extant groups is not necessarily warranted. Using phylogenetic relationships of extant species, we show that five of nine families of North American freshwater fishes exhibit an evolutionary trend of decreasing body size. These trends result from the basal position of large species and the more derived position of small species within families. Such trends may be caused by the invasion of small streams and subsequent isolation and speciation. This pattern, potentially influenced by size-biased dispersal rates and the high percentage of small streams in North America, suggests a scenario that could result in the generation of the size-frequency distribution of North American freshwater fishes.     

Mandibular size and shape variation in the hominins at Dmanisi, Republic of Georgia

The hominin fossils of Dmanisi, Republic of Georgia, present an ideal means of assessing levels of skeletal size and shape variation in a fossil hypodigm belonging to the genus Homo because they have been recovered from a spatially and temporally restricted context. We compare variation in mandible size and shape at Dmanisi to that of extant hominoids and extinct hominins. We use height and breadth measurements of the mandibular corpus at the first molar and the symphysis to assess size, and analyze shape based on size-adjusted (using a geometric mean) versions of these four variables. We compare size and shape variation at Dmanisi relative to all possible pairs of individuals within each comparative taxon using an exact resampling procedure of the ratio of D2600 to D211 and the average Euclidean distance (AED) between D2600 and D211, respectively. Comparisons to extant hominoids were conducted at both the specific and subspecific taxonomic levels and to extinct hominins by adopting both a more, and less speciose, hominin taxonomy. Results indicate that the pattern of variation for the Dmanisi hominins does not resemble that of any living species: they exhibit significantly more size variation when compared to modern humans, and they have significantly more corpus shape variation and size variation in corpus heights and overall mandible size than any extant ape species. When compared to fossil hominins they are also more dimorphic in size (although this result is influenced by the taxonomic hypothesis applied to the hominin fossil record). These results highlight the need to re-examine expectations of levels of sexual dimorphism in members of the genus Homo and to account for marked size and shape variation between D2600 and D211 under the prevailing view of a single hominin species at Dmanisi.     

The Australopithecus assemblage from Sterkfontein Member 4 (South Africa) and the concept of variation in palaeontology

Interpreting morphological variation within the early hominin fossil record is particularly challenging. Apart from the fact that there is no absolute threshold for defining species boundaries in palaeontology, the degree of variation related to sexual dimorphism, temporal depth, geographic variation or ontogeny is difficult to appreciate in a fossil taxon mainly represented by fragmentary specimens, and such variation could easily be conflated with taxonomic diversity. One of the most emblematic examples in paleoanthropology is the Australopithecus assemblage from the Sterkfontein Caves in South Africa. Whereas some studies support the presence of multiple Australopithecus species at Sterkfontein, others explore alternative hypotheses to explain the morphological variation within the hominin assemblage. In this review, I briefly summarize the ongoing debates surrounding the interpretation of morphological variation at Sterkfontein Member 4 before exploring two promising avenues that would deserve specific attention in the future, that is, temporal depth and nonhuman primate diversity.     

A morphometric analysis of extant and early miocene fossil hominoid maxillo-dental specimens

It is demonstrated in this paper that before we can hope to formulate phylogenetic relationships between and amongst fossil hominoid material it is first necessary to sex the material accurately. In order to determine whether the morphological and morphometrical variability seen in fossil specimens is due to sexual or inter species dimorphism, it is necessary to calibrate fossil specimens against extant hominoid species' morphologies. Only after fossil specimens have been sexed is it possible to differentiate between morphologies that are related to sex and those that are species specific. This will help reduce fossil misallocation. A morphometric analysis of extant and fossilProconsul hominoid material is presented. Each fossil specimen has been sexed according to symplesiomorphic sex morphologies as defined in this paper. After the fossil specimens have been sexed they are analyzed using multivariate statistics. The identification of differing sex patterns within the specimens examined here suggests that a new species ofProconsul may have to be considered.     

Metric variation and sexual dimorphism in the dentition of Ouranopithecus macedoniensis

The fossil sample attributed to the late Miocene hominoid taxon Ouranopithecus macedoniensis is characterized by a high degree of dental metric variation. As a result, some researchers support a multiple-species taxonomy for this sample. Other researchers do not think that the sample variation is too great to be accommodated within one species. This study examines variation and sexual dimorphism in mandibular canine and postcanine dental metrics of an Ouranopithecus sample. Bootstrapping (resampling with replacement) of extant hominoid dental metric data is performed to test the hypothesis that the coefficients of variation (CV) and the indices of sexual dimorphism (ISD) of the fossil sample are not significantly different from those of modern great apes. Variation and sexual dimorphism in Ouranopithecus M(1) dimensions were statistically different from those of all extant ape samples; however, most of the dental metrics of Ouranopithecus were neither more variable nor more sexually dimorphic than those of Gorilla and Pongo. Similarly high levels of mandibular molar variation are known to characterize other fossil hominoid species. The Ouranopithecus specimens are morphologically homogeneous and it is probable that all but one specimen included in this study are from a single population. It is unlikely that the sample includes specimens of two sympatric large-bodied hominoid species. For these reasons, a single-species hypothesis is not rejected for the Ouranopithecus macedoniensis material. Correlations between mandibular first molar tooth size dimorphism and body size dimorphism indicate that O. macedoniensis and other extinct hominoids were more sexually size dimorphic than any living great apes, which suggests that social behaviors and life history profiles of these species may have been different from those of living species.     

Extant Lizard Tracks: Variation and Implications for Paleoichnology

In this study, I collected tracks and trackways from nine species of extant lizards representing all five major lizard clades. Previously, tracks from species of only two of these clades were described. Lizard tracks conventionally are regarded as having curved digit imprints that progressively increase in length from digit I to IV, with a smaller digit V directing antero-laterally. However, the zygodactylous feet of chameleons (Calumma parsonii and Furcifer pardalis), the posteriorly directed digit V in the pes of ground-dwelling geckos (Eublepharis macularius) and the rounded feet of blue-tongued skinks (Tiliqua scincoides) did not make “typical” lizard tracks, and demonstrate that even within a limited taxonomic sample there can be considerable variation in the morphologies of lizard tracks. Among the lizards examined, mode of locomotion and how the feet function have more influence on the morphology of tracks than does the phylogenetic affinities of the trackmaker. This preliminary neoichnological study increases the known variation in lizard tracks and aids in interpreting the fossil trackway record by providing comparative information that can be used to identify fossil tracks made by lizards.     

Ontogeny and diachronic changes in sexual dimorphism in the craniofacial skeleton of rhesus macaques from Cayo Santiago, Puerto Rico

Insight into the ontogeny of sexual dimorphism is important to our understanding of life history, ecology, and evolution in primates. This study applied a three-dimensional method, Euclidean Distance Matrix Analysis, to investigate sexual dimorphism and its diachronic changes in rhesus macaque (Macaca mulatta) skulls. Twenty-one landmarks in four functional areas of the craniofacial skeleton were digitized from macaques of known age and sex from the Cayo Santiago collections. Then, a series of mean form matrices, form difference matrices, and growth matrices were computed to demonstrate growth curves, rates and duration of growth, and sexual dimorphism within the neurocranium, basicranium, palate, and face. The inclusion of fully adult animals revealed a full profile of sexual dimorphism. Additionally, we demonstrate for the first time diachronic change in adult sexual dimorphism caused by extended growth in adult females. A quicker growth rate in males from ages 2 to 8 was offset by a longer duration of growth in adult females that resulted in diminished dimorphism between the ages of 8 and 15. Four functional areas showed different sex-specific growth patterns, and the rate and duration of growth in the anterior facial skeleton contributed most to the changing profiles of sexual dimorphism. The late maturation in size of the female facial skeleton corresponds to later and less complete fusion of facial sutures. The prolongation of growth in females is hypothesized to be an evolutionary response to high levels of intrasexual competition, as is found in other primate species such as common chimpanzees with similar colony structure and reproductive behavior. Further investigation is required to determine (1) if this phenomenon observed in craniofacial skeletons is linked to sexual dimorphism in body size, and (2) whether this diachronic change in sexual dimorphism is species specific. The changing profile of sexual dimorphism in adult rhesus macaques suggests caution in studying sexual dimorphism in fossil primate and human forms.     

Growth and sexual dimorphism in a population of hybrid macaques

Growth and sexual dimorphism have long been the focus of investigation for researchers interested in the life history and socioecology of nonhuman primates. Previous research has shown that sex differences in the duration of growth, or bimaturism, are primarily responsible for the sexual dimorphism observed in anthropoid primates with multimale–multifemale social structure, such as macaques. The present study investigates sex differences in patterns of craniofacial and somatometric growth relative to head and body size and relative to dental development in a population of hybrid macaques (Cercopithecidae: Macaca ) from Sulawesi, Indonesia. How these patterns may contribute to sexual dimorphism in this hybrid population is also examined. The results of the study suggest that there is no substantial effect on the levels of sexual dimorphism associated with hybridization in these macaques. Although sex differences in patterns of size-related, or allometric, growth patterns play a significant role in the development of sexual dimorphism for some cranial dimensions in these hybrids, bimaturism seems to be the primary component in the ontogeny of sexual dimorphism in this hybrid population. The observed levels of hybrid dimorphism and the predominant ontogenetic pattern of bimaturism characterized by prolonged male growth are consistent with previously published reports on dimorphism and growth in other cercopithecine primates.     

Sexual dimorphism in the craniofacial growth of the guinea pig (Cavia porcellus)

Variation between the sexes during ontogeny is frequently overlooked in discussions of the phylogenetic patterns of adult sexual dimorphism. Different growth trajectories can produce identical degrees and direction of adult dimorphism and the possibility exists that similarities in adults may be the result of differing growth patterns, suggesting independent evolutionary pathways among species to the seemingly identical adult morphology. We quantified the sexual dimorphism in craniofacial skeletal growth of Cavia porcellus, the guinea pig, using longitudinally collected radiographs. Guinea pigs have male-biased sexual dimorphism in size and in growth parameters, despite literature reports to the contrary. These results, analyzed with equivalent data for five species of rodents, and two outgroups representing similarly sized mammals, a rabbit and a marsupial, indicate that some aspects of sexual differences in growth follow phylogenetic lines, while others are a function of whether the species has male- or female-biased dimorphism.     

Another look at shape variation in the distal femur of Australopithecus afarensis: implications for taxonomic and functional diversity at Hadar

Previous studies have recognized two patterns of distal femoral morphology among the specimens from Hadar (Ethiopia) assigned to Australopithecus afarensis. Size and shape differences between the well-preserved large (AL 333-4) and small (AL 129-1a) distal femora have been used to invoke both taxonomic and functional differences within the A. afarensis hypodigm. Nevertheless, prior studies have not analyzed these specimens in a multivariate context, nor have they compared the pattern of shape differences between the fossils to patterns of sexual dimorphism among extant taxa (i.e., the manner in which males and females differ). This study reexamines morphometric differences between the above specimens in light of observed levels of variation and patterns of sexual dimorphism among extant hominoids. Eight extant reference populations were sampled to provide a standard by which to consider size and shape differences between the fossils. Samples include three populations of modern humans, two subspecies of Pan troglodytes, three subspecies of Gorilla gorilla, Pan paniscus, and Pongo pygmaeus. Using size ratios and scale-free "shape" data (both derived from 2-D coordinate landmarks), size and shape differences between the fossils were evaluated against variation within each reference population using an exact randomization procedure. Growth Difference Matrix Analysis (GDMA) was used to test whether the pattern of morphological differences between the fossils differs significantly from patterns of sexual dimorphism observed among the ten extant groups. Overall morphometric affinities of the fossils to extant taxa were explored using canonical variates analysis (CVA).Results of the randomization tests indicate that the size difference between the Hadar femora can be easily accommodated within most hominoid taxa at the subspecific level (though not within single-sex samples). In addition, the magnitude of shape differences between the fossils can be commonly sampled even within most single-sex samples of a single hominoid subspecies. The pattern of morphological differences between the fossils does not differ statistically from any average pattern of femoral shape dimorphism observed among living hominoids. Moreover, contrary to prior claims, and despite a size disparity between the fossils greater than is typically observed within some chimpanzee and human populations, the two Hadar fossils appear to be much more similar to one another in overall shape than either specimen is to any extant hominoid group.     

Craniodental variation in Paranthropus boisei: a developmental and functional perspective

What levels and patterns of craniodental variation among a fossil hypodigm are necessary to reject the null hypothesis that only a single species is sampled? We suggest how developmental and functional criteria can be used to predict where in the skeleton of fossil hominins we should expect more, or less, within-species variation. We present and test three hypotheses about the factors contributing to craniodental variation in extant primate taxa, and then apply these results to the interpretation of the P. boisei hypodigm. Within the comparative samples of extant Homo, Pan, Gorilla, Pongo, and Colobus, variables from the cranial base, neurocranium, and face that are not subject to high magnitudes of strain have consistently lower levels of intraspecific variation than variables from regions of the face subject to high levels of strain. Dental size variables are intermediate in terms of their reliability. P. boisei is found to have a low degree of variability relative to extant primates for variables shown to be generally useful for testing taxonomic hypotheses. Contrary to the claims of Suwa et al. ([1997] Nature 389:489-492), the recently discovered material from Konso falls within the range of variation of the "pre-Konso" hypodigm of P. boisei for available conventional metrical variables. Those aspects of the Konso material that appear to extend the range of the P. boisei hypodigm involve regions of the skull predicted to be prone to high levels of within-species variation. The approach used in this study focuses on craniodental data, but it is applicable to other regions of the skeleton.     

A simulation test of hominoid species number at Lufeng, China: implications for the use of the coefficient of variation in paleotaxonomy

High dental metric variation in the large hominoid sample from the late Miocene site of Lufeng, China has been interpreted in two ways: (1) there are two morphologically similar species that broadly overlap in size, and (2) there is one species that is more highly sexually dimorphic in dental size, and perhaps in body size, than any extant primate. It has been claimed that the high levels of dental metric variation falsify the single-species hypothesis, which has been viewed implicitly as corroboration of the two-species hypothesis. However, the two-species hypothesis has not been subjected to testing. Here we test the two-species hypothesis using computer simulations to attempt to reproduce the unusual pattern of intrasexual and intersexual dental metric variation observed in the Lufeng postcanine dentition. Conditions of the simulation experiments were optimized to favor the two-species hypothesis. It was found that, although the Lufeng pattern of metric variation could be reproduced by sampling two species, the likelihood of this occurrence was very low even when the conditions were optimized to the point of improbability. We conclude that the likelihood is very high that the Lufeng sample is composed of one species that is more highly sexually dimorphic in the postcanine dentition than any extent primate species. If so, then the high levels of sexual dimorphism and intraspecific dental metric variation in this species violate the central assumption of methods that employ the coefficient of variation (CV) for paleotaxonomy, namely, that neither can lie outside the ranges observed among extant species. Thus, we further conclude that the CV must be used with caution when evaluating the taxonomic composition of fossil samples and, further, that it cannot be used to falsify a single-species hypothesis in any meaningful way. Other fossil hominoid samples with high measures of dental variation may indicate that considerable sexual size dimorphism typified many Eurasian middle–late Miocene hominoids.     

Morphological Integration and the Interpretation of Fossil Hominin Diversity

The fossil record of primate and human evolution cannot provide accurate estimates of within species variation and integration. This means that we cannot directly observe how patterns of integration have evolved over time in this lineage. And yet, our interpretations of fossil diversity are awash with assumptions about variation patterning in precisely these fossil taxa. Most commonly, researchers rely on extant models of variation for interpreting past diversity, by assuming equality of variation (and occasionally covariation) among extant and fossil populations. Yet one of the things we know from studies of integration in primates is that patterns of morphological covariation can differ among even closely related taxa, indicating that they have diverged over evolutionary time, either in response to selection or as the result of neutral evolution. At the same time, overall patterns of integration remain remarkably similar, meaning that in many respects they are highly conserved evolutionarily. Taken together, these seemingly contradictory observations offer an important conceptual framework for interpreting patterns that we observe in the fossil past. This framework dictates that while we can use patterns of covariation in extant taxa as proxies for extinct diversity, and indeed their conserved nature makes them superior to approaches that rely on variation alone, we also need to account for the fact that such patterns change over time, and incorporate that into our models. Here I provide examples using covariation patterns estimated from modern humans and African great apes to demonstrate the extent to which divergence in covariance structure might affect our interpretations of hominin diversity.

Sexual Size Dimorphism,Canine Dimorphism,and Male-Male Competition in Primates

Sexual size dimorphism is generally associated with sexual selection via agonistic male competition in nonhuman primates. These primate models play an important role in understanding the origins and evolution of human behavior. Human size dimorphism is often hypothesized to be associated with high rates of male violence and polygyny. This raises the question of whether human dimorphism and patterns of male violence are inherited from a common ancestor with chimpanzees or are uniquely derived. Here I review patterns of, and causal models for, dimorphism in humans and other primates. While dimorphism in primates is associated with agonistic male mate competition, a variety of factors can affect male and female size, and thereby dimorphism. The causes of human sexual size dimorphism are uncertain, and could involve several non-mutually-exclusive mechanisms, such as mate competition, resource competition, intergroup violence, and female choice. A phylogenetic reconstruction of the evolution of dimorphism, including fossil hominins, indicates that the modern human condition is derived. This suggests that at least some behavioral similarities with Pan associated with dimorphism may have arisen independently, and not directly from a common ancestor.