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.     

Sexual dimorphism in birds: why are there so many different forms of dimorphism?

Variation in the extent of sexual dimorphism among bird species is traditionally attributed to differences in social mating system. However, there are many different forms of dimorphism among birds, and not all of them show an obvious correlation with social mating system. For example, recent work has shown that many highly polygamous species are, in fact, monomorphic, whereas many putatively monogamous species are dimorphic. In this paper we break up sexual dimorphism into subcomponents and then use comparative analyses to examine the pattern of covariation between these subcomponents and various aspects of sexual, social, and parental behaviour. Our first finding is that size dimorphism and plumage-colour dimorphism do not show the same pattern of covariation. Differences in size dimorphism are associated with variation in social mating system and sex differences in parental care, whereas differences in plumage-colour dimorphism are associated with variation in the frequency of extra-bond paternity. These results suggest that size dimorphism is associated with the sort of intrasexual competition described by traditional classifications of social mating system, whereas plumage-colour dimorphism is associated with cryptic female choice. However, when we break up plumage-colour dimorphism according to whether it is due to melanins, carotenoids or structural colours, we find that each category of plumage-colour dimorphism shows a different pattern of covariation. The correlation between overall plumage-colour dimorphism and the rate of extra-bond paternity is due to structural colours, whereas melanin-based dimorphism is associated with sex differences in parental care. The former result is particularly interesting given that new work suggests structural colours are associated with active sexual displays and the reflection of ultraviolet light.     

Sexual selection and canine dimorphism in New World monkeys

Social and ecological factors are important in shaping sexual dimorphism in Anthropoidea, but there is also a tendency for body-size dimorphism and canine dimorphism to increase with increased body size (Rensch's rule) (Rensch: Evolution Above the Species Level. London: Methuen, 1959.) Most ecologist interpret Rensch's rule to be a consequence of social and ecological selective factors that covary with body size, but recent claims have been advanced that dimorphism is principally a consequence of selection for increased body size alone. Here we assess the effects of body size, body-size dimorphism, and social structure on canine dimorphism among platyrrhine monkeys. Platyrrhine species examined are classified into four behavioral groups reflecting the intensity of intermale competition for access to females or to limiting resources. As canine dimorphism increases, so does the level of intermale competition. Those species with monogamous and polyandrous social structures have the lowest canine dimorphism, while those with dominance rank hierarchies of males have the most canine dimorphism. Species with fission-fusion social structures and transitory intermale breeding-season competition fall between these extremes. Among platyrrhines there is a significant positive correlation between body size and canine dimorphism However, within levels of competition, no significant correlation was found between the two. Also, with increased body size, body-size dimorphism tends to increase, and this correlation holds in some cases within competition levels. In an analysis of covariance, once the level of intermale competition is controlled for, neither molar size nor molar-size dimorphism accounts for a significant part of the variance in canine dimorphism. A similar analysis using body weight as a measure of size and dimorphism yields a less clear-cut picture: body weight contributes significantly to the model when the effects of the other factors are controlled. Finally, in a model using head and body length as a measure of size and dimorphism, all factors and the interactions between them are significant. We conclude that intermale competition among platyrrhine species is the most important factor explaining variations in canine dimorphism. The significant effects of size and size dimorphism in some models may be evidence that natural (as opposed to sexual) selection also plays a role in the evolution of increased canine dimorphism.     

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.     

Taxonomic variation in the patterns of craniofacial dimorphism in primates

Understanding sexual dimorphism in living primates is important for interpreting the biological and taxonomic significance of variation in the primate fossil record. In the past two decades, there has been an increasing emphasis on the fact that sexual dimorphism varies in both magnitude and pattern among species. Several studies have suggested that distinct patterns of dimorphism may assist in species recognition and perhaps phylogenetic analysis. This study evaluates patterns of craniofacial dimorphism in samples of 82 anthropoid primates. Dimensions of the viscerocranium tend to be more dimorphic than those of the neurocranium and orbits. Principal components analysis of phylogenetically controlled data demonstrates a basic pattern of dimorphism in overall skull proportions, and a distinction between length and breadth measurements. For any given species there can be substantial variation in the magnitude of dimorphism among dimensions, and different species can show substantially different patterns of dimorphism within and between regions of the skull and jaws. Patterns of dimorphism are clearly associated with phylogeny. Pattern similarity is not dependent on the overall magnitude of craniofacial dimorphism, or body mass dimorphism. Among all anthropoids, there are few combinations of characters that consistently show greater or lesser degrees of dimorphism. Such "stability" of patterns increases within genera. Patterns of dimorphism are likely to be useful for interpreting the taxonomic significance of variation in the fossil record. However, phylogenetic propinquity alone is not reason to use an extant species as a model for variation in an extinct species. Rather, care must be taken to identify stable patterns of dimorphism within a group of closely related extant species.     

Becoming Different But Staying Alike: Patterns of Sexual Size and Shape Dimorphism in Bills of Hummingbirds

Hummingbirds are known for their distinctive patterns of sexual dimorphism, with many species exhibiting sex-related differences in various ecologically-relevant traits, including sex-specific differences in bill shape. It is generally assumed that such patterns are consistent across all hummingbird lineages, yet many taxa remain understudied. In this study we examined patterns of sexual size and sexual shape dimorphism in bills of 32 of 35 species in the monophyletic Mellisugini lineage. We also compared patterns of bill size dimorphism in this group to other hummingbird lineages, using data from 219 hummingbird species. Overall, the presence and degree of sexual size dimorphism was similar across all hummingbird lineages, with the majority of Mellisugini species displaying female-biased sexual size dimorphism, patterns that remain unchanged when analyzed in a phylogenetic context. Surprisingly however, we found that sexual dimorphism in bill shape was nearly absent in the Mellisugini clade, with only 3 of the 32 species examined displaying bill shape dimorphism. Based on observations in other hummingbird lineages, the lack of sexual shape dimorphism in Mellisugini is particularly unusual. We hypothesize that the patterns of sexual size dimorphism observed here may be the consequence of differential selective forces that result from competition for ecological resources. We further propose that an influential mechanism underlying shape dimorphism is competition and niche segregation. Taken together, the evolutionary changes in patterns of sexual shape dimorphism observed in Mellisugini suggest that the evolutionary trends of sexual dimorphism in the Trochilidae are far more dynamic than was previously believed.     

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.     

Intrasexual competition and body weight dimorphism in anthropoid primates

Body weight dimorphism in anthropoid primates has been thought to be a consequence of sexual selection resulting from male-male competition for access to mates. However, while monogamous anthropoids show low degrees of weight dimorphism, as predicted by the sexual selection hypothesis, polygynous anthropoids show high variation in weight dimorphism that is not associated with measures of mating system or sex ratio. This observation has led many to debate the role of other factors such as dietary constraints, predation pressure, substrate constraints, allometric effects, and phylogeny in the evolution of anthropoid weight dimorphism. Here, we re-evaluate variation in adult body weight dimorphism in anthropoids, testing the sexual selection hypothesis using categorical estimates of the degree of male-male intrasexual competition (“competition levels”). We also test the hypotheses that interspecific variation in body weight dimorphism is associated with female body weight and categorical estimates of diet, substrate use, and phylogeny. Weight dimorphism is strongly associated with competition levels, corroborating the sexual selection hypothesis. Weight dimorphism is positively correlated with increasing female body weight, but evidence suggests that the correlation reflects an interaction between overall size and behavior. Arboreal species are, on average, less dimorphic than terrestrial species, while more frugivorous species tend to be more dimorphic than folivorous or insectivorous species. Several alternative hypotheses can explain these latter results. Weight dimorphism is correlated with taxonomy, but so too are competition levels. We suggest that most taxonomic correlations of weight dimorphism represent “phylogenetic niche conservatism”; however, colobines show consistently low degrees of weight dimorphism for reasons that are not clear. Am J Phys Anthropol 103:37–68, 1997. © 1997 Wiley-Liss, Inc.     

Pelvic dimorphism in relation to body size and body size dimorphism in humans

Many mammalian species display sexual dimorphism in the pelvis, where females possess larger dimensions of the obstetric (pelvic) canal than males. This is contrary to the general pattern of body size dimorphism, where males are larger than females. Pelvic dimorphism is often attributed to selection relating to parturition, or as a developmental consequence of secondary sexual differentiation (different allometric growth trajectories of each sex). Among anthropoid primates, species with higher body size dimorphism have higher pelvic dimorphism (in converse directions), which is consistent with an explanation of differential growth trajectories for pelvic dimorphism. This study investigates whether the pattern holds intraspecifically in humans by asking: Do human populations with high body size dimorphism also display high pelvic dimorphism? Previous research demonstrated that in some small-bodied populations, relative pelvic canal size can be larger than in large-bodied populations, while others have suggested that larger-bodied human populations display greater body size dimorphism. Eleven human skeletal samples (total N: male = 229, female = 208) were utilized, representing a range of body sizes and geographical regions. Skeletal measurements of the pelvis and femur were collected and indices of sexual dimorphism for the pelvis and femur were calculated for each sample [ln(M/F)]. Linear regression was used to examine the relationships between indices of pelvic and femoral size dimorphism, and between pelvic dimorphism and female femoral size. Contrary to expectations, the results suggest that pelvic dimorphism in humans is generally not correlated with body size dimorphism or female body size. These results indicate that divergent patterns of dimorphism exist for the pelvis and body size in humans. Implications for the evaluation of the evolution of pelvic dimorphism and rotational childbirth in Homo are considered.     

Intrasexual competition and canine dimorphism in anthropoid primates.

A number of factors, including sexual selection, body weight, body-weight dimorphism, predation, diet, and phylogenetic inertia have been proposed as influences on the evolution of canine dimorphism in anthropoid primates. Although these factors are not mutually exclusive, opinions vary as to which is the most important. The role of sexual selection has been questioned because mating system, which should reflect its strength, poorly predicts variation in canine dimorphism, particularly among polygynous species. Kay et al. (1988) demonstrate that a more refined estimate of intermale competition explains a large proportion of the variation in canine dimorphism in platyrrhine primates. We expand their analysis, developing a more generalized measure of intermale competition based on the frequency and intensity of male-male agonism. We examine the relative influences of predation (inferred by substrate use), female body weight, body-weight dimorphism, diet, and sexual selection on the evolution of anthropoid canine dimorphism. Intermale competition is very strongly associated with canine dimorphism. Predation also has a marked effect on canine dimorphism, in that savanna-dwelling species consistently show greater canine dimorphism than other species, all other factors being held equal. Body-weight dimorphism is also strongly associated with canine dimorphism, though apparently through a common selective basis, rather than through allometric effects. Body weight seems to play only a minor, indirect role in the evolution of canine dimorphism. Diet plays no role. Likewise, we find little evidence that phylogenetic inertia is a constraint on the evolution of canine dimorphism.     

Sexual selection on skeletal shape in Carnivora

Lifetime reproductive success of males is often dependent upon the ability to physically compete for mates. However, species variation in social structure leads to differences in the relative importance of intraspecific aggression. Here, we present a large comparative dataset on sexual dimorphism in skeletal shape in Carnivora to test the hypotheses that carnivorans exhibit sexual dimorphism in skeletal anatomy that is reflective of greater specialization for physical aggression in males relative to females and that this dimorphism is associated with the intensity of sexual selection. We tested these hypotheses using a set of functional indices predicted to improve aggressive performance. Our results indicate that skeletal shape dimorphism is widespread within our sample. Functional traits thought to enhance aggressive performance are more pronounced in males. Phylogenetic model selection suggests that the evolution of this dimorphism is driven by sexual selection, with the best‐fitting model indicating greater dimorphism in polygynous versus nonpolygynous species. Skeletal shape dimorphism is correlated with body size dimorphism, a common indicator of the intensity of male–male competition, but not with mean body size. These results represent the first evidence of sexual dimorphism in the primary locomotor system of a large sample of mammals.     

Relationship of sexual dimorphism in canine size and body size to social,behavioral, and ecological correlates in anthropoid primates

Among anthropoid primates there are interspecific differences in the degree of sexual dimorphism in both body size and canine size. Within the suborder body size dimorphism and canine size dimorphism are positively correlated,r=0.76. This correlation suggests that the two dimorphisms are equally developed in some species, while in other species there is a differential degree of sexual dimorphism. An analysis of these results and their relation to social organization and other ecological variables reveals: (1) the degree of canine size dimorphism is closely related to the amount of male intrasexual selection in a given mating system; and (2) the degree of body size dimorphism is also related to male intrasexual selection, but may be modified (either enhanced or diminished) by selection pressure from factors such as habitat, diet, foraging behavior, antipredator behavior, locomotory behavior, and female preference.     

Big-bodied males help us recognize that females have big pelves

Schultz ([1949] Am. J. Phys. Anthropol. 7:401-424) presented a conundrum: among primates, sexual dimorphism of the pelvis is a developmental adjunct to dimorphism in other aspects of the body, albeit in the converse direction. Among species in which males are larger than females in body size, females are larger than males in some pelvic dimensions; species with little sexual dimorphism in nonpelvic size show little pelvic dimorphism. Obstetrical difficulty does not explain this relationship. The present study addresses this issue, evaluating the relationship between pelvic and femoral sexual dimorphism in 12 anthropoid species. The hypothesis is that species in which males are significantly larger than females in femoral size will have a higher incidence, magnitude, and variability of pelvic sexual dimorphism, with females having relatively larger pelves than males, compared with species monomorphic in femoral size. The results are consistent with the hypothesis. The proposed explanation is that the default pelvic anatomy in adulthood is that of the female; testosterone redirects growth from the default type to that of the male by differentially enhancing and repressing growth among the pelvic dimensions. Testosterone also influences sexual dimorphism of the femur. The magnitude of the pelvic response to testosterone is greater in species that are sexually dimorphic in the femur than in those that are monomorphic.     

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.     

Phylogenetic analyses of dimorphism in primates: evidence for stronger selection on canine size than on body size

Phylogenetic comparative methods were used to analyze the consequences of sexual selection on canine size and canine size dimorphism in primates. Our analyses of previously published body mass and canine size data revealed that the degree of sexual selection is correlated with canine size dimorphism, as well as with canine size in both sexes, in haplorhine but not in strepsirrhine primates. Consistent with these results, male and female canine size was found to be highly correlated in all primates. Since canine dimorphism and canine size in both sexes in haplorhines were found to be not only related to mating system but also to body size and body size dimorphism (characters which are also subject to or the result of sexual selection), it was not apparent whether the degree of canine dimorphism is the result of sexual selection on canine size itself, or whether canine dimorphism is instead a consequence of selection on body size, or vice versa. To distinguish among these possibilities, we conducted matched-pairs analyses on canine size after correcting for the effects of body size. These tests revealed significant effects of sexual selection on relative canine size, indicating that canine size is more important in haplorhine male-male competition than body size. Further analyses showed, however, that it was not possible to detect any evolutionary lag between canine size and body size, or between canine size dimorphism and body size dimorphism. Additional support for the notion of special selection on canine size consisted of allometric relationships in haplorhines between canine size and canine size dimorphism in males, as well as between canine size dimorphism and body size dimorphism. In conclusion, these analyses revealed that the effects of sexual selection on canine size are stronger than those on body size, perhaps indicating that canines are more important than body size in haplorhine male-male competition.     

Sexual dimorphism in cranial shape and size in geomyoid rodents: multivariate and evolutionary perspectives

Geomyoid rodents provide a great study system for the analysis of sexual dimorphism. They are polygynic and many inhabit harsh arid environments thought to promote sexual dimorphism. In fact, there has been extensive work published on the sexual size dimorphism of individual populations and species within this rodent clade. However, little work has been undertaken to assess the evolutionary patterns and processes associated with this sexual dimorphism. We use multivariate analyses of cranial measurements in a phylogenetic framework to determine the distribution of size and shape dimorphism among geomyoids and test for Rensch’s rule. Our results suggest that sexual dimorphism is more common in geomyids than heteromyids, but it is not in fact universal. There is evidence for variation in sexual dimorphism across populations. Additionally, in many taxa, geographic variation appears to overwhelm existing sexual dimorphism. We find support for the repeated independent evolution of shape and size dimorphism across geomyoid taxa, but we do not find support for an association between size and shape dimorphism. There is no evidence for Rensch’s rule in geomyoids, whether at the superfamily or family level. Together, our findings suggest that there is no single explanation for the evolution of sexual dimorphism in geomyoids and that, instead, it is the product of numerous evolutionary events. Future studies incorporating phylogenetic relationships will be necessary to paint a more complete picture of the evolution of sexual dimorphism in geomyoids.     

Sexual dimorphism in the human pelvis: testing a new hypothesis.

Sexual dimorphism in the human pelvis is inferentially related to parturition. Investigators disagree about the identification and obstetric significance of pelvic dimorphism. Benefiting from a large sample of complete skeletons from the Coimbra Identified Skeletal Collection, we show that the dimensions of the true pelvis (birth canal) that are most sexually dimorphic (that is, the dimensions of females are greater than males) are those which are related to biparietal deformation, which often leads to the death of the human neonate. These dimensions are: the anteroposterior diameter of the inlet (index of dimorphism = 108.41), the transverse diameter of the bispinous midplane (index of dimorphism = 117.13) and the transverse diameter of the outlet (index of dimorphism = 112.3). Therefore, sexual dimorphism in the human pelvis is a reflection of differential selection on the two sexes. These results may stimulate further studies with a fresh approach regarding the fossil and comparative evidence for when and how the modern pattern of birth has evolved.     

Evolution of sexual dimorphism in bill size and shape of hermit hummingbirds (Phaethornithinae): a role for ecological causation

Unambiguous examples of ecological causation of sexual dimorphism are rare, and the best evidence involves sexual differences in trophic morphology. We show that moderate female-biased sexual dimorphism in bill curvature is the ancestral condition in hermit hummingbirds (Phaethornithinae), and that it is greatly amplified in species such as Glaucis hirsutus and Phaethornis guy, where bills of females are 60 per cent more curved than bills of males. In contrast, bill curvature dimorphism is lost or reduced in a lineage of short-billed hermit species and in specialist Eutoxeres sicklebill hermits. In the hermits, males tend to be larger than females in the majority of species, although size dimorphism is typically small. Consistent with earlier studies of hummingbird feeding performance, both raw regressions of traits and phylogenetic independent contrasts supported the prediction that dimorphism in bill curvature of hermits is associated with longer bills. Some evidence indicates that differences between sexes of hermit hummingbirds are associated with differences in the use of food plants. We suggest that some hermit hummingbirds provide model organisms for studies of ecological causation of sexual dimorphism because their sexual dimorphism in bill curvature provides a diagnostic clue for the food plants that need to be monitored for studies of sexual differences in resource use.     

Sexual shape dimorphism accelerated by male–male competition,but not prevented by sex-indiscriminate parental care in dung beetles (Scarabaeidae)

Dimorphic sexual differences in shape and body size are called sexual dimorphism and sexual size dimorphism, respectively. The degrees of both dimorphisms are considered to increase with sexual selection, represented by male–male competition. However, the degrees of the two dimorphisms often differ within a species. In some dung beetles, typical sexual shape dimorphisms are seen in male horns and other exaggerated traits, although sexual size dimorphism looks rare. We hypothesized that the evolution of this sexual shape dimorphism without sexual size dimorphism is caused by male–male competition and their crucial and sex-indiscriminate provisioning behaviors, in which parents provide the equivalent size of brood ball with each of both sons and daughters indiscriminately. As a result of individual-based model simulations, we show that parents evolve to provide each of sons and daughters with the optimal amount of resource for a son when parents do not distinguish the sex of offspring and males compete for mates. This result explains why crucial and sex-indiscriminate parental provisioning does not prevent the evolution of sexual shape dimorphism. The model result was supported by empirical data of Scarabaeidae beetles. In some dung beetles, sexual size dimorphism is absent, compared with significant sexual size dimorphism in other horned beetles, although both groups exhibit similar degrees of sexual shape dimorphism in male horns and other exaggerated traits.     

Variability and sexual dimorphism in canine size of Australopithecus and extant hominoids

Among extant hominoids degrees of sexual dimorphism and combined-sex coefficients of variation of canine teeth dimensions are highly correlated. Based on this relationship and coefficients of variation of four species of the genus Australopithecus, we predict degrees of canine dimorphism for these extinct hominids. The estimates show that A. afarensis is as dimorphic as the pygmy chimpanzee, A. boisei slightly less dimorphic than the pygmy chimpanzee, A. robustus slightly more dimorphic than the lar gibbon, while A. africanus overiaps with the lar gibbon as well as a modern human sample. These estimates represent degrees of canine dimorphism substantially lower than results based upon prior sexing of individual specimens. The relationship between canine dimorphism and body weight dimorphism is also analyzed. All four species of Australopithecus are considerably less dimorphic in canine size for their body weight dimorphism than expected. This dissociation of canine size dimorphism and body weight dimorphism is shared with modern humans, and thus represents a unique hominid trait. We interpret the moderate to strong body weight dimorphism in australopithecines as the result of intra- and intersexual selection typical of a polygynous mating structure, while the rather mild canine dimorphism is interpreted in terms of the “developmental crowding” model for reduction in canine size.