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.     

Finger length ratios (2D:4D) in anthropoids implicate reduced prenatal androgens in social bonding

The second‐to‐fourth digit ratio (2D:4D) has been proposed as a biomarker reflecting prenatal androgen effects (PAE), such that individuals with lower ratios have experienced higher PAE than those with higher ratios. 2D:4D has been correlated with a number of sex‐linked traits in humans such as aggression, promiscuity, and competitiveness. In addition, polygynous societies reportedly have lower 2D:4D (higher PAE) than more monogamous populations. This evidence suggests that PAE may be implicated in the development of sexually selected behaviors in humans. To place 2D:4D research into a broader context, we test the relationship between digit ratios and behavior across nonhuman anthropoids; polygynous species, with higher levels of intrasexual competition, should have more pronounced markers of PAE (lower 2D:4D) than pair‐bonded species. Our results accord with those found in humans: 2D:4D is lower in polygynous species and higher (lower PAE) in pair‐bonded species. Old World monkeys have low, and relatively invariant 2D:4D (high PAE), which is coupled with high levels of intrasexual competition. This contrasts with higher and more variable ratios in both great apes and New World monkeys. In addition, both male and female ratios decrease with increasing levels of intrasexual competition. Human ratios are intermediate between pair‐bonded and more promiscuous hominoids. We propose that PAE may be involved in promoting species characteristic social behavior in anthropoids. Am J Phys Anthropol, 2010. © 2009 Wiley‐Liss, Inc.     

Mating systems, sperm competition, and the evolution of sexual dimorphism in birds

Comparative analyses suggest that a variety of factors influence the evolution of sexual dimorphism in birds. We analyzed the relative importance of social mating system and sperm competition to sexual differences in plumage and body size (mass and tail and wing length) of more than 1,000 species of birds from throughout the world. In these analyses we controlled for phylogeny and a variety of ecological and life-history variables. We used testis size (corrected for total body mass) as an index of sperm competition in each species, because testis size is correlated with levels of extrapair paternity and is available for a large number of species. In contrast to recent studies, we found strong and consistent effects of social mating system on most forms of dimorphism. Social mating system strongly influenced dimorphism in plumage, body mass, and wing length and had some effect on dimorphism in tail length. Sexual dimorphism was relatively greater in species with polygynous or lekking than monogamous mating systems. This was true when we used both species and phylogenetically independent contrasts for analysis. Relative testis size was also related positively to dimorphism in tail and wing length, but in most analyses it was a poorer predictor of plumage dimorphism than social mating system. There was no association between relative testis size and mass dimorphism. Geographic region and life history were also associated with the four types of dimorphism, although their influence varied between the different types of dimorphism. Although there is much interest in the effects of sperm competition on sexual dimorphism, we suggest that traditional explanations based on social mating systems are better predictors of dimorphism in birds.     

Size variation in early human mandibles and molars from Klasies River,South Africa: Comparison with other middle and late Pleistocene assemblages and with modern humans

Previous studies of the Middle Stone Age human remains from Klasies River have concluded that they exhibited more sexual dimorphism than extant populations, but these claims have not been assessed statistically. We evaluate these claims by comparing size variation in the best‐represented elements at the site, namely the mandibular corpora and M₂s, to that in samples from three recent human populations using resampling methods. We also examine size variation in these same elements from seven additional middle and late Pleistocene sites: Skhūl, Dolní Věstonice, Sima de los Huesos, Arago, Krapina, Shanidar, and Vindija. Our results demonstrate that size variation in the Klasies assemblage was greater than in recent humans, consistent with arguments that the Klasies people were more dimorphic than living humans. Variation in the Skhūl, Dolní Věstonice, and Sima de los Huesos mandibular samples is also higher than in the recent human samples, indicating that the Klasies sample was not unusual among middle and late Pleistocene hominins. In contrast, the Neandertal samples (Krapina, Shanidar, and Vindija) do not evince relatively high mandibular and molar variation, which may indicate that the level of dimorphism in Neandertals was similar to that observed in extant humans. These results suggest that the reduced levels of dimorphism in Neandertals and living humans may have developed independently, though larger fossil samples are needed to test this hypothesis. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

GEOGRAPHIC VARIATION OF SEXUAL DIMORPHISM IN SIZE OF SAVANNAH SPARROWS (PASSERCULUS SANDWICHENSIS): A TEST OF HYPOTHESES

The Savannah Sparrow (Passerculus sandwichensis) is a widespread and common North American bird that shows both geographic variation and sexual dimorphism in size. I used information from 24 measurements on 1,791 individuals from 51 populations to test two hypotheses (sexual-selection and niche-partitioning) about the evolution of sexual dimorphism. Throughout their range male Savannah Sparrows are larger, on average, than females. This doubtless reflects Darwinian sexual selection, for territorial fights usually involve males, many of whom fail to obtain mates. In some parts of their range, Savannah Sparrows are commonly polygynous, whereas in others they are characteristically monogamous. Among species of American sparrows (subfamily Emberizinae) sexual size dimorphism is generally greater in polygynous species than in monogamous ones. However, I did not find a similar trend among populations of Savannah Sparrows. The amount of dimorphism in all populations of Savannah Sparrows is equivalent in magnitude to that of other species of sparrows that are commonly or regularly polygynous, and it is greater than that of other sparrow species that are characteristically monogamous. The amount of sexual dimorphism, either in overall size or in bill size, does not correlate with species diversity and does not differ between island and mainland populations. These results do not support the niche-variation hypothesis. Size dimorphism is relatively great in populations of Savannah Sparrows that are resident in southwestern salt marshes, and these birds are the only sparrow-like birds that generally breed in these marshes. Dimorphism is, in general, relatively great in marsh-dwelling species in the family Emberizidae. These species are commonly, but not always, polygynous; the mating systems of the salt-marsh Savannah Sparrows are not known. There are no significant differences in the extent of dimorphism among populations of salt-marsh sparrows, and there are few among the non-salt-marsh ones, probably reflecting conservatism in the evolution of size dimorphism.     

Sexual size dimorphism in shorebirds, gulls, and alcids: the influence of sexual and natural selection

Abstract. Charadrii (shorebirds, gulls, and alcids) have an unusual diversity in their sexual size dimorphism, ranging from monomorphism to either male-biased or female-biased dimorphism. We use comparative analyses to investigate whether this variation relates to sexual selection through competition for mates or natural selection through different use of resources by males and females. As predicted by sexual selection theory, we found that in taxa with socially polygynous mating systems, males were relatively larger than females compared with less polygynous species. Furthermore, evolution toward socially polyandrous mating systems was correlated with decreases in relative male size. These patterns depend on the kinds of courtship displays performed by males. In taxa with acrobatic flight displays, males are relatively smaller than in taxa in which courtship involves simple flights or displays from the ground. This result remains significant when the relationship with mating system is controlled statistically, thereby explaining the enigma of why males are often smaller than females in socially monogamous species. We did not find evidence that evolutionary changes in sexual dimorphism relate to niche division on the breeding grounds. In particular, biparental species did not have greater dimorphism in bill lengths than uniparental species, contrary to the hypothesis that selection for ecological divergence on the breeding grounds has been important as a general explanation for patterns of bill dimorphism. Taken together, these results strongly suggest that sexual selection has had a major influence on sexual size dimorphism in Charadrii, whereas divergence in the use of feeding resources while breeding was not supported by our analyses.     

Sexual dimorphism in Laccopithecus robustus, a late miocene hominoid from China

Laccopithecus robustus is a siamang-sized fossil ape from the Miocene site of Lufeng, China. The species is known from a partial cranium, numerous mandibles, and scores of isolated teeth. This species shows striking dental similarities to Pliopithecus from the Miocene of Europe and a number of cranial similarities to extant gibbons. Laccopithecus differs from extant gibbons and resembles other fossil and extant apes in showing marked sexual dimorphism in the size and shape of the canines and anterior lower premolars. Evidence for sexual differences in either the size or shape of other teeth is less clear. There is some evidence for a sexual size dimorphism based on the variability of molar teeth.     

Mating system, sexual dimorphism, and the opportunity for sexual selection in a territorial ungulate

In mammals, species with high sexual size dimorphism tend tohave highly polygynous mating systems associated with high variancein male lifetime reproductive success (LRS), leading to a highopportunity for sexual selection. However, little informationis available for species with weak sexual size dimorphism. Ina long-term study population, we used parentage analysis basedon 21 microsatellite markers to describe, for the first time,variance in male lifetime breeding success (LBS) of roe deer,a territorial ungulate where males weigh less than 10% morethan females. LBS ranged from 0 to 14 (mean = 4.54, variance= 15.5), and its distribution was highly skewed, with only afew males obtaining high LBS and many males failing to breedor siring only one fawn. As predicted for polygynous specieswith low sexual size dimorphism, the standardized variance inmale LBS was low (I_m = 0.75) and was only slightly higher thanthe standardized variance in female LRS (I_f = 0.53), suggestinga low opportunity for sexual selection. The I_m value reportedhere for roe deer is much lower than values reported for highlydimorphic ungulates such as red deer (I_m > 3). We suggestthat, along a continuum of opportunity for sexual selection,roe deer occupy a position closer to monogamous and monomorphicterritorial ungulates than to highly polygynous, sexually dimorphicungulates with dominance rank–based mating systems suchas harems or roving mating systems.     

Evolution of sexual dimorphism in the digit ratio 2D:4D--relationships with body size and microhabitat use in iguanian lizards

The ratio between lengths of digit II and IV (digit ratio 2D:4D) is a morphological feature that likely affects tetrapod locomotor performances in different microhabitats. Modifications of this trait may be triggered by changes in steroids concentrations during embryo development, which might reflect direct selection acting on digit ratio or be solely a consequence of hormonal differences related for example to body size. Here we apply both conventional and phylogenetic analyses on morphological data from 25 lizard species of 3 families of Iguania (Iguanidae, Polychrotidae, and Tropiduridae), in order to verify whether selective pressures related to locomotion in different microhabitats could override the prenatal developmental cues imposed on the digit ratio 2D:4D by differences in body size between males and females. Data suggest that this trait evolved in association with ecological divergence in the species studied, despite the clear effect of body size on the digit ratio 2D:4D. The ecological associations of size-corrected digit ratios were restricted to one sex, and females of species that often use perches exhibited small digit ratios in the front limbs, which translated into larger sexual dimorphism indexes of arboreal species. The results, together with the subsequent discussion, provide outlines for further investigation about possible developmental mechanisms related to the evolution of adaptive changes in digit lengths that may have occurred during the evolution of ecological divergence in squamates.     

Mating system and demographic constraints on the opportunity for sexual selection

In monogamous systems the fitness difference between males due to competition for mates is limited to one female. This constraint presumably impedes the action of sexual selection relative to polygynous systems. In this paper, we use formal selection theory to show how population size and the adult sex ratio constrain the force of sexual selection and phenotypic evolution under monogamy and polygyny. The force of sexual selection is ultimately constrained by the number of males in a population and the theoretical limit to the rate of male phenotypic evolution is realized if a single male mates with one or many females. These results imply that the force of sexual selection is not strictly constrained by monogamy. The constraint on female phenotypic evolution is typically higher than the constraint on males under polygyny and similar to selection on males in monogamous systems. The sexual asymmetry in the force of selection under polygyny--not necessarily weak sexual selection on males of monogamous systems--may explain the prominence of sexual dimorphism in polygynous systems.     

Look in the trees: Hylobatids as evolutionary models for extinct hominins

Studying extant apes is of central importance to paleoanthropology. This approach is informative in inferring how hominin skeletal morphology reflects phylogeny, behavior, development, and ecological context. Traditionally, great apes have dominated the paleoanthropological literature as extant analogs for extinct hominins, to the exclusion of their phylogenetic sister group, the hylobatids. Phylogenetic proximity, large body size, and high encephalization quotients may have contributed to decisions to use great apes as models for hominins. However, if we reexamine hylobatids as extant models for extinct hominins—using modern phylogenetic, behavioral, and ecological data—this clade is uniquely poised to inform future frameworks in paleoanthropology. The following features make hylobatids strong analogs for extinct hominins: taxonomic diversity, the timing of diversification, hybridization between species, small body size, and reduced sexual dimorphism. Based on these shared features, hylobatids offer future opportunities to paleoanthropology, and provide a much richer extant analog than is currently recognized.     

Sexual dimorphism in the face of Australopithecus africanus

Recently discovered crania of Australopithecus africanus from Sterkfontein Member 4 and Makapansgat enlarge the size range of the species and encourage a reappraisal of both the degree and pattern of sexual dimorphism. Resampling methodology (bootstrapping) is used here to establish that A. africanus has a greater craniofacial size range than chimpanzees or modern humans, a range which is best attributed to a moderately high degree of sexual dimorphism. Compared to other fossil hominins, this variation is similar to that of Homo habilis (sensu lato) but less than that of A. boisei. The finding of moderately high dimorphism is corroborated by a CV-based estimate and ratios between those specimens considered to be male and those considered to be female. Inferences about the pattern of craniofacial dimorphism in the A. africanus face currently rely on the relationship of morphology and size. Larger specimens, particularly Stw 505, show prominent superciliary eminences and glabellar regions, but in features related in part to canine size, such as the curvature of the infraorbital surface, large and small specimens of A. africanus are similar. In this respect, the pattern resembles that of modern humans more so than chimpanzees or lowland gorillas. A. africanus may also show novel patterns of sexual dimorphism when compared to extant hominines, such as in the form of the anterior pillar. However, males of the species do not exhibit characteristics of more derived hominins, such as A. robustus. Am J Phys Anthropol 108:97–127, 1999. © 1999 Wiley-Liss, Inc.     

The origins of sexual dimorphism in body size in ungulates

Jarman (1974) proposed a series of relationships between habitat use, food dispersion, and social behavior and hypothesized a series of evolutionary steps leading to sexual dimorphism in body size through sexual selection in African antelope species. The hypothesis states that sexual size dimorphism evolved in a three-step process. Initially, ancestral monomorphic and monogamous ungulate species occupying closed habitats radiated into open grassland habitats. Polygynous mating systems then rapidly evolved in response to the aggregation of males and females, perhaps in relation to the clumped distribution of food resources in open habitats. Subsequently, size dimorphism evolved in those species occupying open habitats, but not in species that remained in closed habitats or retained monogamy. This hypothesis has played an important role in explaining the origins of sexual dimorphism in mammals. However, the temporal sequence of the events that Jarman proposed has never been demonstrated. Here we use a phylogeny of extant ungulate species, along with maximum-likelihood statistical techniques, to provide a test of Jarman's hypothesis.     

Sexual dimorphism,mating system,and effect of phylogeny in De Brazza's monkey (Cercopithecus neglectus)

De Brazza's monkey (Cercopithecus neglectus), like other guenons, shows marked sexual dimorphism in an array of features. While strong sexual dimorphism is generally associated with a polygynous mating system, populations of De Brazza's monkeys in Gabon are reportedly monogamous. An explanation of this unique phenomenon is offered here. Patterns of sexual dimorphism are examined for morphology, growth and development, behavior, and ecology, and field and captive studies on the social organization and mating system of De Brazza's monkey and congeneric guenon species are reviewed. Based on the findings, it is postulated that 1) De Brazza's monkeys are not strictly monogamous, but exhibit interpopulational variation in their mating system, from facultative monogamy to mild polygyny; 2) marked sexual dimorphism most likely reflects the effect of the historical-phylogenetic factor; ie, it represents a holdover of a degree of dimorphism established earlier in evolutionary history when the degree of polygyny Was higher; and 3) lessening in the degree of polygyny and a tendency toward monogamy represents a consequence of selection toward small group size. Small group size, a unique antipredator strategy, and failure to form polyspecific associations are ultimately most likely the result of intragroup and interspecific competition and predation pressure.     

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.     

Does Social Mating System Influence Nest Defence Behaviour in Great Reed Warbler (Acrocephalus arundinaceus) Males?

In birds with biparental care, males and females often conflict over how much care to provide to their offspring and it may be substantially influenced by increased level of polygamy. In accordance with sexual conflict theory, males of socially polygynous bird species provide much less care to their nestlings than do males of most socially monogamous species. Most of previous studies, however, have used feeding behaviour as an index for variations in male parental care only. However, this may be skewed if polygynous males compensate for lower feeding assistance through the provision of other parental care such as protection of nests from predators. In this paper, we examine nest defence behaviour in the facultatively polygynous great reed warbler with respect to sex and type of social mating system. We recorded latency to the first arrival, distance from the predator and defensive reaction of each parent towards a human intruder. Socially polygynous males with two simultaneously active nests defended primary females’ nests less vigorously than socially monogamous males, whereas no differences were found between monogamous and primary females. Generally, however, they took a bigger role in nest defence than males in all cases. Our results support an idea that sexual conflict is driven by polygamy and that type of social mating system can influence nest defence behaviour of facultatively polygynous birds. This finding should be taken into consideration when studying nest defence parental care in polygynous mating systems.     

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.     

Implications of Male and Female Contributions to Sexual Size Dimorphism for Inferring Behavior in the Hominin Fossil Record

Sexual dimorphism is commonly used to directly infer or support reconstructions of social behavior in early hominins. This is often done by comparing the magnitude of sexual size dimorphism to that seen in extant primates and extrapolating a likely social behavior. Such comparisons are of limited value, though, allowing only the inference of strong male–male competition when dimorphism is strong. Recent studies have begun to focus on the selective factors that impact female body size, and thereby size dimorphism. Considerations of changes in male and female size in the fossil record potentially allow insight into the meaning of changes in sexual dimorphism through time. To illustrate, I compare estimates of body mass dimorphism for four hominin taxa to assess changes in male and female size. Assuming that early Homo represents a single taxon, sexual size dimorphism increased in early Homo through an increase in male size, but was subsequently reduced through an increase in female size in Homo erectus. This would imply a significant increase in sexual selection acting on males in early Homo. An increase in female size with a loss of dimorphism in Homo erectus would imply a simultaneous shift in female optimal body size through selection for increased female fecundity, and/or an increase in female resource abundance, coupled with a shift in selection acting on male size. Although none of these inferences are certain, the exercise illustrates the potential for considering how dimorphism changes through time, rather than simply focusing on the magnitude of size dimorphism in isolation.     

Canine tooth size variability in primates

I present an analysis of canine tooth size variability in male and female primates. The coefficient of variation (CV = SD X 100/mean) as an index of canine size variability proved to be dependent on mean canine size in males and, to a lower extent, in females. Therefore, variability tends to increase with increasing values of mean canine size. Using residuals from the regression of log SD on log mean canine size in male and female primates, I analysed the contribution of diet, habitat and mating system to canine size variability. Habitat and mating system are known to influence to a certain extent the degree of sexual dimorphism in canine size. Given the well-known relationship between sexual dimorphism and phenotypic variability, it was suggested that these factors might influence variability in canine size. Everything else being equal, males of polygynous species are characterized by more variable canine sizes than males of monogamous species. Habitat and diet did not contribute to the level of variability observed in either males or females. It is proposed that a high level of variability in canine size may be related to the likelihood that enlarged canines evolved as a result of male-male competition for mates in polygynous species.