Adult proportionality in small-bodied foragers: a test of ecogeographic expectations

If predictable, ecogeographic patterning in body size and proportions of human populations can provide valuable information regarding human biology, adaptation to local environments, migration histories, and health, now and in the past. This paper evaluates the assumption that small-bodied Later Stone Age (LSA) foragers of Southern Africa show the adult proportions that would be expected of warm-adapted populations. Comparisons are also made with small-bodied foragers from the Andaman Islands (AI). Indices including brachial, crural, limb element length to skeletal trunk height, and femoral head and bi-iliac breadth to femoral length were calculated from samples of LSA (n = 124) and AI (n = 31) adult skeletons. Samples derived from the literature include those from high (Europe), middle (North Africa), and low (Sub-Saharan Africa) latitude regions. The LSA and AI samples match some but not all expected ecogeographic patterns for their particular regions of long term habitation. For most limb length to skeletal trunk height indices the LSA and AI are most similar to the other mid-latitude sample (North Africans). However, both groups are similar to low latitude groups in their narrow bi-iliac breadths, and the AI display relatively long radii. Proportions of LSA and AI samples also differ from those of African pygmies. In regions like southern-most Africa, that do not experience climatic extremes of temperature or humidity, or where small body size exists through drift or selection, body size, and proportions may also be influenced by nonclimatic variables, such as energetic efficiency.     

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.     

An exploration of adult body shape and limb proportions at Kellis 2, Dakhleh Oasis,Egypt

Several studies have shown that the human body generally conforms to the ecogeographical expectations of Bergmann's and Allen's rules; however, recent evidence suggests that these expectations may not hold completely for some populations. Egypt is located at the crossroads of Sub‐Saharan Africa, Southern Europe, and the Near East, and gene flow among groups in these regions may confound ecogeographical patterning. In this study, we test the fit of the adult physique of a large sample (N = 163) of females and males from the Kellis 2 cemetery (Dakhleh Oasis, Egypt) against ecogeographical predictions. Body shape (i.e., body mass relative to stature) was assessed by the femur head diameter to bicondylar femur length index (FHD/BFL), and brachial and crural indices were calculated to examine intralimb proportions. Body shape in the Kellis 2 sample is not significantly different from high‐latitude groups and a Lower Nubian sample, and intralimb proportions are not significantly different from mid‐latitude and other low‐latitude groups. This study demonstrates the potential uniqueness of body shape and intralimb proportions in an ancient Egyptian sample, and further highlights the complex relationship between ecogeographic patterning and adaptation. Am J Phys Anthropol 153:496–505, 2014. © 2013 Wiley Periodicals, Inc.     

Effects of parity on pelvic size and shape dimorphism in Mus

The pelvis is a sexually dimorphic structure and although the causes of that dimorphism have long been studied, relatively little is known regarding the effects of partuitive events on the magnitude of that dimorphism. Here, we use a sample of Mus musculus domesticus to contrast dimorphism in body length and os coxae size and shape between males and parous and nulliparous females. We also test for correlations between relative litter size (L/M) and relative offspring size (O/M) with body length and os coxae size and shape in parous females. Males had greater body length than nulliparous females but were not different from parous females. Females as a whole had the largest os coxae, with parous females having the largest and males the smallest. Os coxae shape was also significantly different between groups and was most divergent between parous females and males than between nulliparous females and males. Os coxae shape differences between females are associated with differences in body length between females and O/M is correlated with os coxae shape in parous females such that females with the largest offspring have the most divergent shapes along the relative warp one axis. Pelvic shape differences between males and females were consistent with previous findings in other taxa which identify the pubo‐ischial complex as the primary region of dimorphism. J. Morphol. 2009. © 2009 Wiley‐Liss, Inc.     

Body proportions in ancient Andeans from high and low altitudes

Living human populations from high altitudes in the Andes exhibit relatively short limbs compared with neighboring groups from lower elevations as adaptations to cold climates characteristic of high-altitude environments. This study compares relative limb lengths and proportions in pre-Contact human skeletons from different altitudes to test whether ecogeographic variation also existed in Andean prehistory. Maximum lengths of the humerus, radius, femur, and tibia, and femoral head breadth are measured in sex-specific groups of adult human skeletons (N = 346) from the central (n = 80) and the south-central (n = 123) Andean coasts, the Atacama Desert at 2,500 m (n = 102), and the southern Peruvian highlands at 2,000-3,800 m (n = 41). To test whether limb lengths vary with altitude, comparisons are made of intralimb proportions, limb lengths against body mass estimates derived from published equations, limb lengths against the geometric mean of all measurements, and principal component analysis. Intralimb proportions do not statistically differ between coastal groups and those from the Atacama Desert, whereas intralimb proportions are significantly shorter in the Peruvian highland sample. Overall body size and limb lengths relative to body size vary along an altitudinal gradient, with larger individuals from coastal environments and smaller individuals with relatively longer limbs for their size from higher elevations. Ecogeographic variation in relation to climate explains the variation in intralimb proportions, and dietary variation may explain the altitudinal cline in body size and limb lengths relative to body size. The potential effects of gene flow on variation in body proportions in Andean prehistory are also explored.     

Allometry of head and body size in Holocene foragers of the South African Cape

Opportunities to assess morphological allometry in small-bodied human populations are rare. The foragers of the Later Stone Age of the South African Cape are characteristically small-bodied. Previous studies have shown that during the period of ca. 3500 to 2000 years BP (uncalibrated (14) C dates), the regional population shows transient reduced stature, body mass, and cranial size, a pattern that has been tentatively tied to demographic pressure on resources. This study examines the relationships among cranial size (centroid size) and body size (femoral length, femoral head diameter, and bi-iliac breadth) during the second half of the Holocene (N = 62). Reduced major axis regression indicates negative allometry of cranial centroid size with body size. Residuals (from ordinary least squares regression of cranial centroid size on body size) are regressed on radiocarbon date to examine temporal changes in the relationship between cranial and body size. Cranial and pelvic sizes are most conserved through time, while more ancient skeletons possess shorter femora and smaller femoral heads. The relationship between cranial centroid size and femoral length shows larger and more variable residuals at more recent dates, indicating a greater or more variable disassociation between cranial size and stature relative to more ancient skeletons. A similar, but nonsignificant relationship exists between cranial size and bi-iliac breadth. These results provide insights into the use of aspects of body size and proportionality in the assessment of health in past populations.     

Testicle size of orang-utans in relation to body size

Few data are available for assessing the relative testicle size of orang-utans, Pongo pygmaeus, so measures were obtained for 31 individuals of varying age. It was shown that the volume of the testicles, calculated from in situ measures of testicle length and breadth, closely approximates testicle weight when multiplied by the specific gravity of solid tissue. Growth curves for body weight and data published for wild specimens were evaluated to obtain the weight most characteristic of male Pongo, and the ratio of testicle weight to body weight was calculated. The mean ratio for individuals with fully adult stature is 0.034, similar to but smaller than that of humans at about 0.050, and larger than the ratios reported for 5 gorillas at 0.013. The testicles mature faster than the body, however, so the mean ratio for young adult orang-utans is about 0.056 and resembles the ratio for humans more closely than the full adults. The differences between the ratios for a monogamous gibbon species, orang-utans, and humans is accounted for when testicle size relative to the weight of the female is considered. This is consistent with a sperm dilution effect produced by variation in the size of the female reproductive tract. The small relative testicle size of the gorilla is anomalous and requires verification as does the application of female size to scale the testicles. © 1993 Wiley-Liss, Inc.     

Protection of obstetric dimensions in a small-bodied human sample

In human females, the bony pelvis must find a balance between being small (narrow) for efficient bipedal locomotion, and being large to accommodate a relatively large newborn. It has been shown that within a given population, taller/larger-bodied women have larger pelvic canals. This study investigates whether in a population where small body size is the norm, pelvic geometry (size and shape), on average, shows accommodation to protect the obstetric canal. Osteometric data were collected from the pelves, femora, and clavicles (body size indicators) of adult skeletons representing a range of adult body size. Samples include Holocene Later Stone Age (LSA) foragers from southern Africa (n = 28 females, 31 males), Portuguese from the Coimbra-identified skeletal collection (CISC) (n = 40 females, 40 males) and European-Americans from the Hamann-Todd osteological collection (H-T) (n = 40 females, 40 males). Patterns of sexual dimorphism are similar in the samples. Univariate and multivariate analyses of raw and Mosimann shape-variables indicate that compared to the CISC and H-T females, the LSA females have relatively large midplane and outlet canal planes (particularly posterior and A-P lengths). The LSA males also follow this pattern, although with absolutely smaller pelves in multivariate space. The CISC females, who have equally small stature, but larger body mass, do not show the same type of pelvic canal size and shape accommodation. The results suggest that adaptive allometric modeling in at least some small-bodied populations protects the obstetric canal. These findings support the use of population-specific attributes in the clinical evaluation of obstetric risk.     

Airway thermal volume in humans and its relation to body size

Serikov, Vladimir B., E. Heidi Jerome, Neal W. Fleming,Peter G. Moore, Frederick A. Stawitcke, and Norman C. Staub.Airway thermal volume in humans and its relation to body size.J. Appl. Physiol. 83(2): 668-676, 1997.The objective of this study was to investigate the influence ofvolume ventilation(E) andcardiac output () on the temperature of the expiredgas at the distal end of the endotracheal tube in anesthetized humans.In 63 mechanically ventilated adults, we used a step decrease in thehumidity of inspired gas to cool the lungs. After change from humid todry gas ventilation, the temperature of the expired gas decreased. Weevaluated the relationship between the inverse monoexponential timeconstant of the temperature fall (1/) and eitherE or . WhenE wasincreased from 5.67 ± 1.28 to 7.14 ± 1.60 (SD) l/min(P = 0.02), 1/ did not changesignificantly [from 1.25 ± 0.38 to 1.21 ± 0.51 min¹,P = 0.81]. In the 11 patients in whom changed during the study period(from 5.07 ± 1.81 to 7.38 ± 2.45 l/min,P = 0.02), 1/ increasedcorrespondingly from 0.89 ± 0.22 to 1.52 ± 0.44 min¹(P = 0.003). We calculated the airwaythermal volume (ATV) as the ratio of the measured values to 1/ and related it to the body height (BH):ATV (liters) = 0.086 BH (cm)  9.55 (r = 0.90).

     

Morphological variability of the shape of striped red mullet Mullus surmuletus in relation to stock discrimination between the Bay of Biscay and the eastern English Channel

Truss analysis and length measurements were made on 168 striped red mullet Mullus surmuletus. Multivariate statistical analyses with principal component analysis and partial redundancy analysis (pRDA) were used on these measurements to evaluate the influence of maturity, sex and geographical area distribution on body shape. Truss measurements were important to quantify and discriminate changing body shape, presumably due to changing environmental conditions. Sexual dimorphism was not observed and juveniles could be distinguished from adults based on their body shape. More importantly, M. surmuletus occurring in different geographical areas could be differentiated using this method. Based on pRDA, a significant difference of head morphological dimensions was observed between populations occurring in the eastern English Channel and those occurring in the Bay of Biscay, suggesting that fish from these areas could represent two subpopulations.     

Optimal diving behaviour for foraging in relation to body size

Overall, large animals dive longer and deeper than small animals; however, after the difference in body size is taken into account, smaller divers often tend to make relatively longer dives. Neither physiological nor theoretical explanations have been provided for this paradox. This paper develops an optimal foraging diving model to demonstrate the effect of body size on diving behaviour, and discusses optimal diving behaviour in relation to body size. The general features of the results are: (1) smaller divers should rely more heavily on anaerobic respiration, (2) larger divers should not always make longer dives than smaller divers, and (3) an optimal body size exists for each diving depth. These results explain the relatively greater diving ability observed in smaller divers, and suggest that if the vertical distribution of prey in the water column is patchy, there is opportunity for a population of diving animals to occupy habitat niches related to body size.     

Unveiling the environmental drivers of intraspecific body size variation in terrestrial vertebrates

Aim

Whether intraspecific spatial patterns in body size are generalizable across species remains contentious, as well as the mechanisms underlying these patterns. Here we test several hypotheses explaining within-species body size variation in terrestrial vertebrates including the heat balance, seasonality, resource availability and water conservation hypotheses for ectotherms, and the heat conservation, heat dissipation, starvation resistance and resource availability hypotheses for endotherms.

Location

Global.

Time period

1970–2016.

Major taxa studied

Amphibians, reptiles, birds and mammals.

Methods

We collected 235,905 body size records for 2,229 species (amphibians = 36; reptiles = 81; birds = 1,545; mammals = 567) and performed a phylogenetic meta-analysis of intraspecific correlations between body size and environmental variables. We further tested whether correlations differ between migratory and non-migratory bird and mammal species, and between thermoregulating and thermoconforming ectotherms.

Results

For bird species, smaller intraspecific body size was associated with higher mean and maximum temperatures and lower resource seasonality. Size–environment relationships followed a similar pattern in resident and migratory birds, but the effect of resource availability on body size was slightly positive only for non-migratory birds. For mammals, we found that intraspecific body size was smaller with lower resource availability and seasonality, with this pattern being more evident in sedentary than migratory species. No clear size–environment relationships were found for reptiles and amphibians.

Main conclusions

Within-species body size variation across endotherms is explained by disparate underlying mechanisms for birds and mammals. Heat conservation (Bergmann's rule) and heat dissipation are the dominant processes explaining biogeographic intraspecific body size variation in birds, whereas in mammals, body size clines are mostly explained by the starvation resistance and resource availability hypotheses. Our findings contribute to a better understanding of the mechanisms behind species adaptations to the environment across their geographic distributions.     

Habitat primary production and the evolution of body size within the hartebeest clade

Local adaptation is a key process in the evolution of biological diversity but relatively few studies have identified the selective forces that drive trait divergence at low taxonomic levels, particularly amongst mammals. Variation in body size across taxa is fundamental as shown by allometric relationships with numerous physiological, morphological and life-history traits. Differences in adult size across cohorts within populations of temperate ungulates are determined by variation in trophic resource availability during growth, suggesting that natural selection might promote the evolution of size divergence across sister taxa through local adaptation to variation in habitat productivity. We tested this hypothesis in the hartebeest ( Alcelaphu s sp.), an antelope lineage including eight extant (or recently extinct) allopatric subspecies that evolved within the last million years and colonized all the African savannahs. We predicted that body size across the subspecies should correlate positively with habitat productivity across taxon ranges. Mean body size of all the hartebeest taxa was quantified using skull length from museum specimens, and climatic variables were used as surrogates of habitat productivity. Body size across subspecies was positively correlated with rainfall, suggesting that variation in habitat primary production may drive morphological evolution between taxa. Focusing at a low taxonomic level has allowed us to identify a critical selective force that may shape divergence in body size, without the confounding effect of variation in trophic niche. © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 92 , 431–440.     

Length of activity season drives geographic variation in body size of a widely distributed lizard

Understanding the factors that drive geographic variation in life history is an important challenge in evolutionary ecology. Here, we analyze what predicts geographic variation in life‐history traits of the common lizard, Zootoca vivipara, which has the globally largest distribution range of all terrestrial reptile species. Variation in body size was predicted by differences in the length of activity season, while we found no effects of environmental temperature per se. Females experiencing relatively short activity season mature at a larger size and remain larger on average than females in populations with relatively long activity seasons. Interpopulation variation in fecundity was largely explained by mean body size of females and reproductive mode, with viviparous populations having larger clutch size than oviparous populations. Finally, body size‐fecundity relationship differs between viviparous and oviparous populations, with relatively lower reproductive investment for a given body size in oviparous populations. While the phylogenetic signal was weak overall, the patterns of variation showed spatial effects, perhaps reflecting genetic divergence or geographic variation in additional biotic and abiotic factors. Our findings emphasize that time constraints imposed by the environment rather than ambient temperature play a major role in shaping life histories in the common lizard. This might be attributed to the fact that lizards can attain their preferred body temperature via behavioral thermoregulation across different thermal environments. Length of activity season, defining the maximum time available for lizards to maintain optimal performance, is thus the main environmental factor constraining growth rate and annual rates of mortality. Our results suggest that this factor may partly explain variation in the extent to which different taxa follow ecogeographic rules.     

Fusion of coccyx to sacrum in humans: prevalence, correlates, and effect on pelvic size, with obstetrical and evolutionary implications

Humans do not have a tail, but we have four rudimentary coccygeal vertebrae. This study considers several issues pertaining to fusion of the coccyx to the sacrum, including prevalence, sexual differences, effect on pelvic size, and obstetrical and evolutionary implications. Previous research on sacral-coccygeal fusion has reported: (1) lower prevalence in females than males, (2) prevalence increases with age, (3) range in prevalence among 13 samples from 0 to 72%, and (4) obstetrical complications. This study uses a sample of 2,354 American skeletons of known sex, age 20 years and older to ascertain prevalence of sacral-coccygeal fusion and to evaluate some of its correlates. Results show that the sexes do not differ in prevalence of sacral-coccygeal fusion for five of seven decades of life, but that prevalence does increase with advancing age-from 24 to 47% from the third to eighth decades of life in females. Pelvimetric analysis of 132 females shows that those with sacral-coccygeal fusion have a shorter posterior sagittal diameter of the outlet compared to those without fusion; more than half of those with sacral-coccygeal fusion have an obstetrically contracted posterior sagittal diameter. Shortening of the posterior sagittal diameter is important, because its conjoint occurrence with a narrow subpubic arch may result in an obstetrically inadequate outlet. This study concludes that sacral-coccygeal fusion is a principal contributor to the evolution of sexual dimorphism in sacral angulation, which is a determinant of the length of the posterior sagittal diameter of the outlet.     

Autosomal variation for male body size and sperm competition phenotypes is uncorrelated in Drosophila melanogaster

Correlations between male body size and phenotypes impacting post-copulatory sexual selection are commonly observed during the manipulation of male body size by environmental rearing conditions. Here, we control for environmental influences and test for genetic correlations between natural variation in male body size and phenotypes affecting post-copulatory sexual selection in Drosophila melanogaster. Dry weights of virgin males from 90 second-chromosome and 88 third-chromosome substitution lines were measured. Highly significant line effects (p<0.001) documented a genetic basis to variation in male body size. No significant correlations were identified between male body size and the components of sperm competitive ability. These results suggest that natural autosomal variation for male body size has little impact on post-copulatory sexual selection. If genetic correlations exist between male body size and post-copulatory sexual selection then variation in the sex chromosomes are likely candidates, as might be expected if sexually antagonistic coevolution was responsible.