Polymorphic aspects of male anthropoid honing premolars.

A recent study has indicated that heritage may be an important component in explanations of interspecies variation in male anthropoid maxillary canines. To further test this hypothesis, comparative data for the maxillary canine's occlusal partner, the honing mandibular premolar, are presented. Relative to body mass, measurements of the honing premolar--length of the honing facet, mesiodistal length, and buccolingual breadth and shape (as expressed by the ratio of length to breadth)--show a similar but stronger heritage component than measurements of the maxillary canine. Results indicate that canine variation among male anthropoids has not yet been explained and that quantitative analyses of this issue will necessarily require a methodological approach which incorporates heritage as a factor.     

Canine size and shape in male anthropoid primates

While studies of canine dimorphism in primates are common, only a few have examined canine tooth size independently within each sex. Recently, Greenfield and Washburn (Am. J. Phys. Anthropol. 84:17–34, 1991) proposed that there are two types of male canines which reflect different allometric scaling patterns of canine crown height against canine occlusal dimensions. They also suggest that proportional canine shape, measured as canine crown height (or projection) relative to the occlusal dimensions, provides an estimate of the canine's function as a weapon, though they provide no test of this hypothesis. This analysis critically examines the claim that there are two types of male canines among anthropoids. It then tests the hypothesis that relative male canine size (measured against body weight) and proportional canine shape are related to estimates of intermale competition, diet, and substrate (used as a surrogate measure of predation pressure). While there is strong taxonomic variation in canine size and shape among male anthropoids, no evidence is found for two discrete canine types. Rather, within families and subfamilies, canine dimensions scale isometrically against body weight and against each other, with a continuum of canine shapes among different taxa. While variation in male canine size is associated with intermale competition and substrate, even when taxonomic variation is controlled, variation in proportional canine shape is not. Neither canine size nor shape are generally associated with variation in diet. © 1993 Wiley-Liss, Inc.     

Canine size, shape, and bending strength in primates and carnivores

Anthropoid primates are well known for their highly sexually dimorphic canine teeth, with males possessing canines that are up to 400% taller than those of females. Primate canine dimorphism has been extensively documented, with a consensus that large male primate canines serve as weapons for intrasexual competition, and some evidence that large female canines in some species may likewise function as weapons. However, apart from speculation that very tall male canines may be relatively weak and that seed predators have strong canines, the functional significance of primate canine shape has not been explored. Because carnivore canine shape and size are associated with killing style, this group provides a useful comparative baseline for primates. We evaluate primate maxillary canine tooth size, shape and relative bending strength against body size, skull size, and behavioral and demographic measures of male competition and sexual selection, and compare them to those of carnivores. We demonstrate that, relative to skull length and body mass, primate male canines are on average as large as or larger than those of similar sized carnivores. The range of primate female canine sizes embraces that of carnivores. Male and female primate canines are generally as strong as or stronger than those of carnivores. Although we find that seed-eating primates have relatively strong canines, we find no clear relationship between male primate canine strength and demographic or behavioral estimates of male competition or sexual selection, in spite of a strong relationship between these measures and canine crown height. This suggests either that most primate canines are selected to be very strong regardless of variation in behavior, or that primate canine shape is inherently strong enough to accommodate changes in crown height without compromising canine function.     

The influence of body proportions on femoral and tibial midshaft shape in hunter‐gatherers

Variation in femoral and tibial diaphyseal shape is used as an indicator of adaptation to patterns of terrestrial mobility. Recent experimentation has implied that lower limb diaphyseal shape may be primarily influenced by lower limb length, and less so by mobility patterns. If valid, this would, at most, render previous interpretations of mobility patterns based on analyses of diaphyseal shape questionable, and, at least, require additional standardization that considers the influence of limb length. Although the consequences could be profound, this implication has yet to be directly tested. Additionally, the influence of body breadth on tibial shape (and to a lesser extent femoral shape) remains uncertain. Tibial and femoral cross‐sectional midshaft shape measurements, taken from nine Pleistocene and Holocene skeletal populations, were compared against lower limb length, limb segment length, and bi‐iliac breadth. Generally, limb length and limb segment length do not significantly influence femoral or tibial midshaft shape. After controlling for body mass greater bi‐iliac breadth is associated with a relative mediolateral strengthening of the femoral midshaft, while the influence of a wider body shape (BIB/length) is associated with a relative M‐L strengthening of the tibia and femur of males, and the tibia of females. We conclude that; (1) mechanical interpretations of lower limb diaphyseal shape are most parsimonious due to the lack of evidence for a consistent relationship between segment length and shape; however, (2) further work is required to investigate the influence of bi‐iliac breadth on both femoral and tibial midshaft shape. Am J Phys Anthropol, 2010. © 2010 Wiley‐Liss, Inc.     

Intrasexual selection in Verreaux's sifaka (Propithecus verreauxi verreauxi)

Studies of sexual selection show that both female choice and male-male competition can influence the evolution and expression of male phenotypes. In this regard, it is important to determine the functional basis through which male traits influence variation in male reproductive success. In this study, we estimate the strength and type of sexual selection acting on adult males in a population of wild lemur, Verreaux's sifaka (Propithecus verreauxi verreauxi). The data used in this study were collected at Beza Mahafaly Special Reserve, southwest Madagascar. We conducted paternity analyses on 70 males in order to estimate the distribution of reproductive success in this population. Paternity data were combined with morphometric data in order to determine which morphological traits covary with male fitness. Five morphological traits were defined in this analysis: body size, canine size, torso shape, arm shape, and leg shape. We utilized phenotypic selection models in order to determine the strength and type of selection acting directly on each trait. Our results show that directional selection acts on leg shape (a trait that is functionally related to locomotor performance), stabilizing selection acts on body mass and torso shape, and negative correlational selection acts on body mass and leg shape. We draw from biomechanical and kinematic studies of sifaka locomotion to provide a functional context for how these traits influence male mating competition within an arboreal environment. Verreaux's sifaka and many other gregarious lemurs are sexually monomorphic in body mass and canine size, despite a high frequency and intensity of male-male aggressive competition. Our results provide some insight into this paradox: in our population, there is no directional selection acting on body mass or canine size in males. The total pattern of selection implicates that behaviors relating to locomotor performance are more important than behaviors relating to fighting ability during intrasexual contests.     

Effect of composition on shape of bird eggs

Numerous studies over the past 90 years have described the various bird egg shapes in mathematical terms but few studies have considered the underlying reasons for such interspecific egg shape variability. This study investigated how the size and composition, i.e. proportions of shell, yolk and albumen, were associated with egg shape. Geometric morphometrics were used to generate principal components, which were analysed in relation to taxonomy (i.e. avian order) and degree of neonatal developmental maturity, which correlates with egg composition. The analysis confirmed previous results that most of the variation in shape is associated with degree of elongation (i.e. length divided by breadth) and asymmetry (i.e. position of the broadest part of the egg away from the mid‐point of the egg's length). Egg shape reflected avian order but not developmental maturity. The degree of elongation of an egg is related to absolute egg mass and the proportion of yolk. By contrast, the degree of asymmetry is related to the proportion of shell and the mass of the egg relative to female body mass. Although significant, the models explained little of the variation in egg shape and so it was concluded that other factors, such as pelvis size and shape, could be more important in determining egg shape in birds.     

Ontogenetic and individual variation in size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis

The present study investigates relationships among size, shape and speed in the Australian agamid lizard Amphibolurus nuchalis . Maximal running speed, body mass, snout-vent length, tail length, fore- and hind limb spans and thigh muscle mass were measured in 68 field-fresh individuals spanning the entire ontogenetic size range (1.3 48 g). Relative lengths of both foreand hind limbs decrease with increasing body mass (= negative allometry), whereas relative tail length and thigh muscle mass increase with body mass (= positive allometry). Repeatable and significant differences in maximal running speed exist among individuals. Maximal running speed scales as (body mass)^0.161, and 59% of the variation in maximal speed was related to body mass. Based on the results of the present and previous studies, data on scaling of body proportions alone appear inadequate to infer scaling relationships of functional characters such as top speed.
Surprisingly, individual variation in maximal speed is not related to individual variation in shape (relative limb, tail and body lengths). These components of overall shape are not independent; individuals tended to have either relatively long or relatively short limbs, tails and bodies for their body mass. Even the significant difference in multivariate shape between adult males and females has no measurable consequences for maximal speed. Speeds of field-fresh animals did not vary on a seasonal basis, and eight weeks of captivity had no effect on maximal running speeds. Gravid females and long-term (obese) captive lizards were both approximately 12% slower than field-fresh lizards.     

Endocranial cast of Hexian Homo erectus from South China

In this paper, we present data on the morphological features and linear measurements for the Hexian Homo erectus and other comparative endocasts, in order to highlight variation during human brain evolution. The endocast of Hexian was reconstructed in 1982, and an endocranial volume of 1,025 ml was estimated. The geological age is about 412 ka, or roughly contemporaneous with the Zhoukoudian (ZKD) specimens. There are some differences between Hexian and the modern Chinese male endocasts in our sample, including low position of the greatest breadth, low maximum height, a well-marked and prominent frontal keel, the flat surface of the frontal lobes, prominent sagittal keel along the center frontal and parietal lobes, depressed Sylvian areas and parietal lobes superiorly, strong posterior projection of the occipital lobes, anterior position of the cerebellar lobes relative to the occipital lobes, and the relative simplicity of the meningeal vessels. Compared with the ZKD, Indonesian, and African Homo erectus specimens, Hexian has more morphological features in common with ZKD. Principal component analyses indicate that Hexian is closest to the ZKD Homo erectus compared with the modern Chinese and other Homo erectus, but its great breadth distinguishes it. Metric analyses show that the brain height, frontal breadth, cerebral height, frontal height, and parietal chord from Homo erectus to modern humans increased, while the length, breadth, frontal chord, and occipital breadth did not change substantially.     

Calculating egg volume when shape differs: when are equations appropriate?

ABSTRACT Egg volume is often calculated from length and breadth, assuming little variation in egg shape within species. However, egg shape may vary among females, over time, or within clutches. If this variation is not predictable, volume should be estimated using another method. For example, in some species, egg shape changes consistently with laying order and volume can be estimated using laying‐order specific equations. We measured the length, breadth, volume, and mass of 249 Magellanic Penguin (Spheniscus magellanicus) eggs and regressed volume on length and breadth for first and second eggs separately. Differences between measured and calculated volume averaged <1% for both eggs. Using linear measurements, we calculated egg volume for 7085 clutches from 1983 to 2006. First and second eggs had similar calculated volumes (P > 0.05) in 19 of 24 yr; when they differed, the second egg was larger. In contrast, using single equations typically employed to compute egg volume, we erroneously concluded that first eggs were usually (20 of 24 yr) significantly larger than second eggs. Our regression equations should be applicable to other Spheniscus penguins because ratios of elongation coefficients (length divided by breadth) of first and second eggs are similar among these species. In other species of birds where egg shape changes with laying order, the volume of a sample of eggs should be measured to develop regression equations specific to the species and laying order. Slight variation in egg volume can have important evolutionary and ecological implications that would be erroneously interpreted without field tests to measure volume.     

Body size and body shape in early hominins - implications of the Gona Pelvis

Discovery of the first complete Early Pleistocene hominin pelvis, Gona BSN49/P27, attributed to Homo erectus, raises a number of issues regarding early hominin body size and shape variation. Here, acetabular breadth, femoral head breadth, and body mass calculated from femoral head breadth are compared in 37 early hominin (6.0-0.26 Ma) specimens, including BSN49/P27. Acetabular and estimated femoral head sizes in the Gona specimen fall close to the means for non-Homo specimens (Orrorin tugenesis, Australopithecus africanus, Paranthropus robustus), and well below the ranges of all previously described Early and Middle Pleistocene Homo specimens. The Gona specimen has an estimated body mass of 33.2 kg, close to the mean for the non-Homo sample (34.1 kg, range 24-51.5 kg, n = 19) and far outside the range for any previously known Homo specimen (mean = 70.5 kg; range 52-82 kg, n = 17). Inclusion of the Gona specimen within H. erectus increases inferred sexual dimorphism in body mass in this taxon to a level greater than that observed here for any other hominin taxon, and increases variation in body mass within H. erectus females to a level much greater than that observed for any living primate species. This raises questions regarding the taxonomic attribution of the Gona specimen. When considered within the context of overall variation in body breadth among early hominins, the mediolaterally very wide Gona pelvis fits within the distribution of other lower latitude Early and Middle Pleistocene specimens, and below that of higher latitude specimens. Thus, ecogeographic variation in body breadth was present among earlier hominins as it is in living humans. The increased M-L pelvic breadth in all earlier hominins relative to modern humans is related to an increase in ellipticity of the birth canal, possibly as a result of a non-rotational birth mechanism that was common to both australopithecines and archaic Homo.     

Convergent evolution of sexual shape dimorphism in Diptera

Several patterns of sexual shape dimorphism, such as male body elongation, eye stalks, or extensions of the exoskeleton, have evolved repeatedly in the true flies (Diptera). Although these dimorphisms may have evolved in response to sexual selection on male body shape, conserved genetic factors may have contributed to this convergent evolution, resulting in stronger phenotypic convergence than might be expected from functional requirements alone. I compared phenotypic variation in body shape in two distantly related species exhibiting sexually dimorphic body elongation: Prochyliza xanthostoma (Piophilidae) and Telostylinus angusticollis (Neriidae). Although sexual selection appears to act differently on male body shape in these species, they exhibited strikingly similar patterns of sexual dimorphism. Likewise, patterns of within-sex shape variation were similar in the two species, particularly in males: relative elongation of the male head capsule, antenna, and legs was associated with reduced head capsule width and wing length, but was nearly independent of variation in thorax length. However, the two species presented contrasting patterns of static allometry: male sexual traits exhibited elevated allometric slopes in T. angusticollis, but not in P. xanthostoma. These results suggest that a shared pattern of covariation among traits may have channeled the evolution of sexually dimorphic body elongation in these species. Nonetheless, static allometries may have been shaped by species-specific selection pressures or genetic architectures.     

Sexual dimorphism in postcranial skeletal shape suggests male-biased specialization for physical competition in anthropoid primates

Sexual dimorphism often arises as a response to selection on traits that improve a male's ability to physically compete for access to mates. In primates, sexual dimorphism in body mass and canine size is more common in species with intense male–male competition. However, in addition to these traits, other musculoskeletal adaptations may improve male fighting performance. Postcranial traits that increase strength, agility, and maneuverability may also be under selection. To test the hypothesis that males, as compared to females, are more specialized for physical competition in their postcranial anatomy, we compared sex-specific skeletal shape using a set of functional indices predicted to improve fighting performance. Across species, we found significant sexual dimorphism in a subset of these indices, indicating the presence of skeletal shape sexual dimorphism in our sample of anthropoid primates. Mean skeletal shape sexual dimorphism was positively correlated with sexual dimorphism in body size, an indicator of the intensity of male–male competition, even when controlling for both body mass and phylogenetic relatedness. These results suggest that selection on male fighting ability has played a role in the evolution of postcranial sexual dimorphism in primates.     

Genetic variation in egg dimensions in natural populations of the Great Tit

Length and breadth of eggs were measured in ringed populations of the Great Tit. During a part of the study volume and weight were also measured, but this did not give additional information, viz. variation in specific weight of fresh eggs and deviations from calculated volume were within the limits of precision. Only in small eggs are length and breadth positively correlated.In two populations, a major part (60–80%) of the variation in the clutch means of egg length, egg breadth, shape index and egg volume is only found between clutches of different females. The absence of correlation between different female partners of one male and the similarity of female repeatability to heritability estimates based on daughter-mother regression lead to the conclusion that 60–80% of the variation in egg dimensions is genetic.The implications for a potential rapid response to selection resulting in a micro-evolutionary change are discussed.     

Determinants of local abundance in a major radiation of Australian passerines (Aves: Meliphagoidea)

Aim  To identify the factors that contribute to variation in abundance (population density), and to investigate whether habitat breadth and diet breadth predict macroecological patterns in a suborder of passerine birds (Meliphagoidea).
Location  Australia (including Tasmania).
Methods  Mean abundance data were collated from site surveys of bird abundance (the Australian Bird Count); range size and latitudinal position data from published distribution maps; and body mass and diet breadth information from published accounts. A diversity index of habitats used (habitat breadth) was calculated from the bird census data. We used bivariate correlation and multiple regression techniques, employing two phylogenetic comparative methods: phylogenetic generalized least squares and independent contrasts.
Results  Body mass and latitude were the only strong predictors of abundance, with larger-bodied and lower-latitude species existing at lower densities. Together, however, body mass and latitude explained only 11.1% of the variation in mean abundance. Range size and habitat breadth were positively correlated, as were diet breadth and body mass. However, neither range size, nor habitat breadth and diet breadth, explained patterns in abundance either directly or indirectly.
Main conclusions  Levels of abundance (population density) in meliphagoid birds are most closely linked to body mass and latitudinal position, but not range size. As with many other macroecological analyses, we find little evidence for aspects of niche breadth having an effect on patterns of abundance. We hypothesize that evolutionary age may also have a determining effect on why species tend to be rarer (less abundant) in the tropics.     

Intrapopulation variation in stature and body proportions: social status and sex differences in an Italian medieval population (Trino Vercellese, VC)

The phenotypic expression of adult body size and shape results from synergistic interactions between hereditary factors and environmental conditions experienced during growth. Variation in body size and shape occurs even in genetically relatively homogeneous groups, due to different occurrence, duration, and timing of growth insults. Understanding the causes and patterns of intrapopulation variation can foster meaningful information on early life conditions in living and past populations. This study assesses the pattern of biological variation in body size and shape attributable to sex and social status in a medieval Italian population. The sample includes 52 (20 female, 32 male) adult individuals from the medieval population of Trino Vercellese, Italy. Differences in element size and overall body size (skeletal height and body mass) were assessed through Monte Carlo methods, while univariate non-parametric tests and Principal Component Analysis (PCA) were employed to examine segmental and overall body proportions. Discriminant Analysis was employed to determine the predictive value of individual skeletal elements for social status in the population. Our results highlight a distinct pattern in body size and shape variation in relation to status and sex. Male subsamples exhibit significant postcranial variation in body size, while female subsamples express smaller, nonsignificant differences. The analysis of segmental proportions highlighted differences in trunk/lower limb proportions between different status samples, and PCA indicated that in terms of purely morphological variation high status males were distinct from all other groups. The pattern observed likely resulted from a combination of biological factors and cultural practices.     

Body size, body shape, and long bone strength in modern humans

To identify behaviorally significant differences in bone structure it is first necessary to control for the effects of body size and body shape. Here the scaling of cross-sectional geometric properties of long bone diaphyses with different "size" measures (bone length, body mass, and the product of bone length and body mass) are compared in two modern human populations with very different body proportions: Pecos Pueblo Amerindians and East Africans. All five major long bones (excluding the fibula) were examined. Mechanical predictions are that cortical area (axial strength) should scale with body mass, while section modulus (bending/torsional strength) should scale with the product of body mass and moment arm length. These predictions are borne out for section moduli, when moment arm length is taken to be proportional to bone length, except in the proximal femoral diaphysis, where moment arm length is proportional to mediolateral body breadth (as would be expected given the predominance of M-L bending loads in this region). Mechanical scaling of long bone bending/torsional strength is similar in the upper and lower limbs despite the fact that the upper limb is not weight-bearing. Results for cortical area are more variable, possibly due to a less direct dependence on mechanical factors. Use of unadjusted bone length alone as a "size" measure produces misleading results when body shape varies significantly, as is the case between many modern and fossil hominid samples. In such cases a correction factor for body shape should be incorporated into any "size" standardization.     

New model for estimating the relationship between surface area and volume in the human body using skeletal remains

A new model for estimating human body surface area and body volume/mass from standard skeletal metrics is presented. This model is then tested against both 1) “independently estimated” body surface areas and “independently estimated” body volume/mass (both derived from anthropometric data) and 2) the cylindrical model of Ruff. The model is found to be more accurate in estimating both body surface area and body volume/mass than the cylindrical model, but it is more accurate in estimating body surface area than it is for estimating body volume/mass (as reflected by the standard error of the estimate when “independently estimated” surface area or volume/mass is regressed on estimates derived from the present model). Two practical applications of the model are tested. In the first test, the relative contribution of the limbs versus the trunk to the body's volume and surface area is compared between “heat-adapted” and “cold-adapted” populations. As expected, the “cold-adapted” group has significantly more of its body surface area and volume in its trunk than does the “heat-adapted” group. In the second test, we evaluate the effect of variation in bi-iliac breadth, elongated or foreshortened limbs, and differences in crural index on the body's surface area to volume ratio (SA:V). Results indicate that the effects of bi-iliac breadth on SA:V are substantial, while those of limb lengths and (especially) the crural index are minor, which suggests that factors other than surface area relative to volume are driving morphological variation and ecogeographical patterning in limb prorportions. Am J Phys Anthropol 156:614–624, 2015. © 2014 Wiley Periodicals, Inc.     

How Humans Differ from Other Animals in Their Levels of Morphological Variation

Animal species come in many shapes and sizes, as do the individuals and populations that make up each species. To us, humans might seem to show particularly high levels of morphological variation, but perhaps this perception is simply based on enhanced recognition of individual conspecifics relative to individual heterospecifics. We here more objectively ask how humans compare to other animals in terms of body size variation. We quantitatively compare levels of variation in body length (height) and mass within and among 99 human populations and 848 animal populations (210 species). We find that humans show low levels of within-population body height variation in comparison to body length variation in other animals. Humans do not, however, show distinctive levels of within-population body mass variation, nor of among-population body height or mass variation. These results are consistent with the idea that natural and sexual selection have reduced human height variation within populations, while maintaining it among populations. We therefore hypothesize that humans have evolved on a rugged adaptive landscape with strong selection for body height optima that differ among locations.     

Take-off flight performance in the butterfly Pararge aegeria relative to sex and morphology: a quantitative genetic assessment

Adult fitness components may strongly depend on variation in locomotory performance such as flight; this variation can be sex specific. Fast take-off to intercept females and competing males is an essential behavioral component of the territorial perching behavior in male speckled wood butterflies (Pararge aegeria L.). Females on the other hand avoid frequent take-offs particularly under suboptimal temperatures, typically showing fewer but longer flights than males. We estimated the heritability of take-off acceleration performance under suboptimal body temperatures by a restricted maximum-likelihood model. We calculated genetic correlations between this performance and a selection of morphological traits: size (body mass), flight muscle investment (relative thorax mass), and wing shape (forewing aspect ratio). Our results show significant additive genetic variation for mean acceleration performance and a similar but nonsignificant trend (P= 0.08) for maximal acceleration performance during take-off in males (h(2)= 0.15). In females, heritability was not significantly different from zero for either of the acceleration performance measures. Morphological traits and take-off performance were genetically linked in a sex-specific way. In males, relative thorax mass and forewing aspect ratio were positively genetically correlated with acceleration performance. In females, there was a negative genetic correlation between acceleration performance and abdomen mass, but not with residual abdomen mass (i.e., regressed on total body mass). To fully understand the evolution of sexual differences in flight performances and morphology, several other flight performances will have to be included. This multifunctional nature of flight and its consequences for the evolutionary study of flight has not yet been fully appreciated in the literature.     

COPULATORY BEHAVIOR,GENITAL MORPHOLOGY,AND MALE FERTILIZATION SUCCESS IN WATER STRIDERS

Recent theoretical and empirical interest in postmating processes have generated a need for increasing our understanding of the sources of variance in fertilization success among males. Of particular importance is whether such postmating sexual selection merely reinforces the effects of premating sexual selection or whether other types of male traits are involved. In the current study, we document large intraspecific variation in last male sperm precedence in the water strider Gerris lateralis. Male relative paternity success was repeatable across replicate females, showing that males differ consistently in their ability to achieve fertilizations. By analyzing shape variation in male genital morphology, we were able to demonstrate that the shape of male intromittent genitalia was related to relative paternity success. This is the first direct experimental support for the suggestion that male genitalia evolve by postmating sexual selection. A detailed analysis revealed that different components of male genitalia had different effects, some affecting male ability to achieve sperm precedence and others affecting male ability to avoid sperm precedence by subsequent males. Further, the effects of the shape of the male genitalia on paternity success was in part dependent on female morphology, suggesting that selection on male genitalia will depend on the frequency distribution of female phenotypes. We failed to find any effects of other morphological traits, such as male body size or the degree of asymmetry in leg length, on fertilization success. Although males differed consistently in their copulatory behavior, copulation duration was the only behavioral trait that had any significant effect on paternity.