Morphological variation and sexual dimorphism of the cephalic scales in Lacerta bilineata

The Western green lizard (Lacerta bilineata) is a lacertid distributed throughout Mediterranean and Central Europe. Little is known about the morphological variability and sexual shape differences in this species. In this paper, the variation of the cephalic scales in L. bilineata is analysed by means of a geometric morphometric approach. A main structural pattern is characterized by negative allometry of the frontal and interparietal areas, and positive allometry of the parietal and frontoparietal scales. Sexual differences are described both in size and shape. In males, the scales are generally larger, with relative shortening of the frontal area, frontoparietal enlargement, occipital lengthening and bulging of the parietal scales, which compresses and narrows the interparietal and occipital areas midsagittally. This pattern is based on a shared allometric trajectory, with males displaying a peramorphic morphotype. However, males show some shape differences in the occipital area that are not size‐related, and cannot be interpreted in terms of general head enlargement. This structural trajectory can be related to the development of the skull, but the role of soft tissues (temporal, nuchal and masticatory muscles) must also be considered. The development of the jaw and nuchal muscles involved in intra‐ and intersexual behaviours could have played a pivotal role in the evolution of this pattern.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. I. Single trait heritabilities

The use of skeletal nonmetric traits in studies of biological relationships often involves the assumption that variation in these traits is genetic. Studies of nonmetric traits in human groups and in inbred strains of mice and rabbits have indicated a genetic component to nonmetric trait variation. Skeletons of animals with known matrilineage membership were obtained from the Cayo Santiago skeletal collection in order to obtain a direct estimate of the heritabilities of several nonmetric traits in the free-ranging population of rhesus macaques on Cayo Santiago. Falconer's (1965) method was used to calculate heritability. Heritability estimates range from zero to one, and half of them are greater than 0.5. This indicates that there is a considerable amount of genetic variation for these traits among the Cayo macaques. There is a significant tendency for traits scoring the number of foramina to have lower heritabilities than those scoring hyperstotic or hypostotic traits.     

Bone mineral density, osteopenia, and osteoporosis in the rhesus macaques of Cayo Santiago

This cross-sectional study investigates metabolic bone disease and the relationship between age and bone mineral density (BMD) in males and females of a large, well-documented skeletal population of free-ranging rhesus monkeys (Macaca mulatta), from the Caribbean Primate Research Center Museum collection from Cayo Santiago, Puerto Rico. The sample consists of 254 individuals aged 1.0-20+ years. The data consist of measurements of bone mineral content and bone mineral density, obtained from dual-energy X-ray absorptiometry (DEXA), of the last lumbar vertebra from each monkey. The pattern of BMD differs between male and female rhesus macaques. Females exhibit an initial increase in BMD with age, with peak bone density occurring around age 9.5 years, and remaining constant until 17.2 years, after which there is a steady decline in BMD. Males acquire bone mass at a faster rate, and attain a higher peak BMD at an earlier age than do females, at around 7 years of age, and BMD remains relatively constant between ages 7-18.5 years. After age 7 there is no apparent effect of age on BMD in the males of this sample; males older than 18.5 years were excluded due to the presence of vertebral osteophytosis, which interferes with DEXA. The combined frequency of osteopenia and osteoporosis in this population is 12.4%. BMD values of monkeys with vertebral wedge fractures are generally higher than those of virtually all of the nonfractured osteopenic/osteoporotic individuals, thus supporting the view that BMD as measured by DEXA is a useful but imperfect predictor of fracture risk, and that low BMD may not always precede fractures in vertebral bones. Other factors such as bone quality (i.e., trabecular connectivity) should also be considered. The skeletal integrity of a vertebra may be compromised by the loss of key trabeculae, resulting in structural failure, but the spine may still show a BMD value within normal limits, or within the range of osteopenia.     

The relationship between cranial metric and nonmetric traits in the rhesus macaques from Cayo Santiago

This study addresses the relationship between cranial metric variables and nonmetric traits using the skeletal sample of rhesus macaques from Cayo Santiago. Discriminant function analysis is used to study the metric differences between macaque crania grouped according to the presence or absence of nine nonmetric traits. The computation of total structure coefficients from the discriminant function analyses provides information regarding how closely each metric variable is related to the discriminant functions derived. Total structure coefficients have not been used previously in the study of the relationship between metric and nonmetric traits. The results of the analysis are interpreted using an explicit approach to cranial morphogenesis-functional cranial analysis. It is concluded that the relationship between cranial metric and nonmetric traits is explicable in terms of a common developmental pathway shared by the two types of traits. Identification of the specific etiology of nonmetric traits depends on future anatomical studies or organisms throughout the period of nonmetric trait development.     

Reproductive performance, population dynamics and anthropometrics of the free-ranging Cayo Santiago rhesus macaques

This report summarizes demographic data collected on the Cayo Santiago colony of rhesus monkeys from 1976-1983 and compares the results with those from 1959-1964 [8,9]. For males and nonpregnant/nonlactating, pregnant, and lactating females mean (+/- 1 SD), body weights, crown-rump lengths, and ponderal indices are tabulated for each age on a large (n = 586) single sampling of this free-ranging population of macaques.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques on Cayo Santiago. II. Phenotypic,genetic, and environmental correlations between traits

The general lack of phenotypic correlation among skeletal nonmetric traits has been interpreted as indicating a lack of genetic correlation among these traits. Nonmetric traits scored on animals in the skeletal collection of rhesus macaques from Cayo Santiago are used to calculate phenotypic, genetic, and environmental correlations between traits. The results show that even when phenotypic correlations are low, there may be large, significant genetic correlations among these traits. The genetic correlation pattern suggests that genes which affect nonmetric trait variation act primarily at a local level in the cranium, even though there are genes with pleiotropic effects on skeletal nonmetric traits throughout the cranium. Environmental and phenotypic correlations do not show this neighborhood pattern of correlation.     

Power and limits of DNA-profiling in primate populations: Paternity assessment in rhesus macaques from Cayo Santiago

Paternity testing was performed in one social group (S) of rhesus macaques from Cayo Santiago, Puerto Rico. In 11/15 cases, sires could be identified comparing the multilocus DNA profiles of 19 males to those of the corresponding mother/child dyads. All 19 males could be excluded from paternity in the remaining four cases. Decision making was partly based on likelihoods of DNA profiles, and the theoretical model underlying these calculation is described. In a second social group (M), held in captivity, paternity testing was impeded by a deficit of maternal bands and by an increased extent of band sharing of mothers and their infants. Some possible explanations for these findings, including increased homozygosity in group M, are discussed.     

Quantitative genetics of skeletal nonmetric traits in the rhesus macaques of Cayo Santiago. III. Relative heritability of skeletal nonmetric and metric traits

This study addresses the long-standing controversy in skeletal biology concerning the relative utility of skeletal metric and nonmetric traits for studies of biological relationship. This controversy centers on the relative heritability of these two trait sets. This paper presents heritabilities for a series of skeletal metric and nonmetric traits measured with the same sample of mother-offspring pairs from the Cayo Santiago skeletal collection of rhesus macaques. Skeletal nonmetric traits display significantly greater heritability estimates than metric traits. This difference is due primarily to the high heritability estimates of hyperostotic nonmetric traits. Foraminal traits are not significantly more heritable than skeletal metric traits. The generality of this pattern of heritability values, in which hyperostotic nonmetric traits are more highly heritable than foraminal nonmetric and metric traits, depends on future empirical study of the correlation of heritability values in populations and theoretical work.     

Fusion patterns of craniofacial sutures in rhesus monkey skulls of known age and sex from Cayo Santiago

Bones of the face and cranial vault meet at sutural boundaries. These sutures are of great importance for craniofacial growth. Although the effects that the sutures have on modulating craniofacial strains have been investigated, how sutural fusion influences primate craniofacial biomechanics and adaptation are less considered. Confounding this problem is the lack of any systematic data on patterns of craniofacial sutural fusion from animals of known age and sex. This study examined the status of 28 sutures in Macaca mulatta skulls from a collection of animals of known age and sex from Cayo Santiago, Puerto Rico. Survival analysis showed that most animals died before all sutures fused. There was high variation in the age at which individual sutures or sutural sections were fused in M. mulatta, and significant differences in the amount of sutural fusion among regions and between males and females. Intensive fusion of sutures took place between ages 5 and 15. Sutures in the facial area tended to be less fused than in the cranial vault. Between adolescence and adulthood, males tended to have more sutural fusion than females, especially in the facial area. These differences might be biomechanical adaptations during ontogeny to craniofacial sexual dimorphism. These findings enrich our understanding of variation in sutural morphology in rhesus monkeys. Comparative information across primate species is essential for understanding the biomechanics of craniofacial form throughout primate evolution.     

Ontogeny of craniofacial sexual dimorphism in the orangutan (Pongo pygmaeus). I: face and palate

The orangutan is widely recognized as a highly dimorphic species. An ontogenetic approach to the study of sexual dimorphism can assist researchers in understanding both where and when these differences develop. In this study, 357 orangutans from Borneo were divided into five developmental stages representing infancy to mature adulthood. Three-dimensional (3D) coordinate data from 16 landmarks representing the face and palate were analyzed by means of a Euclidean distance matrix analysis (EDMA), a quantitative method for the comparison of forms. Three separate analyses (an age-specific static comparison of forms, a sex-specific analysis of growth trajectories, and an intersex comparison of patterns of relative growth) were carried out with the intent to describe the rate, timing, magnitude, and pattern of growth in the orangutan face and palate. The results indicate that generally males and females share a similar, but not identical, pattern of growth or local form change, but differ in growth rate, timing, and magnitude of difference. Dimorphism in the face and palate can be localized in infancy and traced throughout all age intervals. Orangutan females grow slightly faster than males from infancy to adolescence, at which time male growth exceeds female growth. Female growth ceases with the advent of adulthood, while male growth continues (i.e., both the number and magnitude of the dimorphic dimensions increase). Males and females are similar in facial dimensions and growth related to the orbits, upper face, and palate width. They maintain these similarities throughout development. However, they differ in facial and nasal height, palate length, snout projection, depth of the nasopharynx, and hafting of the face onto the skull. The face broadens and the zygomatic bone flares dramatically in adult males, corresponding to the development of cheek pads. While growth patterns are similar between the two sexes, they differ in the lateral orbit, snout projection, and hafting of the face onto the cranium. Adult dimorphism is the result of growth patterns experienced throughout life, and it is not equally expressed across the cranium. An understanding of patterns of dimorphism, along with the magnitude of difference, may be helpful for interpreting dimorphism in the fossil record.     

The hemogram,serum biochemistry,and electrolyte profile of the free-ranging Cayo Santiago rhesus macaques (Macaca mulatta)

Sixty-four male and 33 female free-ranging rhesus monkeys (Macaca mulatta) from one of six social groups on the island of Cayo Santiago, Puerto Rico, were surveyed to establish normal values for the hemogram and serum biochemical and electrolytes for the colony. Mean values (± 1 SD) are reported by sex for each of three age groups (2–3, 4–9 and ≥ 10 years). All adult females (≥ 4 years) were pregnant. There were significant differences for a number of variables compared to the range reported in the existing literature, and among the age and sex groups in the sampled population.     

Osteoarthritis of the hand in nonhuman primates: A clinical,radiographic, and skeletal survey of Cayo Santiago rhesus macaques

Abstract: We describe the relative prevalence and pattern of distribution of osteoarthritis (OA) in the hands of elderly (>15 years) rhesus macaques using clinical, radiographic, and skeletal examinations. In the clinical study the prevalence of nodes was 72% and 16% in the distal inter-phalangeal joints (DIPJ) and proximal inter-phalangeal joints (PIPJ), respectively, 31% of all monkeys had polyarticular nodes. Radiographic OA was present in 55%, 9.1%, and 0% of the DIPJs, PIPJs, and thumb bases, respectively. Skeletal OA as defined by joint surface eburnation for the DIPJ, PIPJ, and thumb base were 16%, 8%, and 2%, respectively. A similar pattern of hand OA with humans is described except for the thumb base OA. This may be due to the relatively rudimentary manipulative role of the macaque thumb. The finding of polyarticular nodal OA raises the possibility of a common pathogenensis for IPJ OA amongst primates.     

Ontogeny and sex alter the effect of predation on body shape in a livebearing fish: sexual dimorphism, parallelism, and costs of reproduction

Predation can cause morphological divergence among populations, while ontogeny and sex often determine much of morphological diversity among individuals. We used geometric morphometrics to characterize body shape in the livebearing fish Brachyrhaphis rhabdophora to test for interactions between these three major shape-determining factors. We assessed shape variation between juveniles and adults of both sexes, and among adults for populations from high- and low-predation areas. Shape differed significantly between predation regimes for all juveniles regardless of sex. As males grew and matured into adults, ontogenetic shape trajectories were parallel, thus maintaining shape differences in adult males between predation environments. However, shape of adult females between predation environments followed a different pattern. As females grew and matured, ontogenetic shape trajectories converged so that shape differences were less pronounced between mature females in predator and nonpredator environments. Convergence in female body shape may indicate a trade-off between optimal shape for predator evasion versus shape required for the livebearing mode of reproduction.     

Ontogenic changes and sexual dimorphism in the head, mouth and teeth of the lesser spotted dogfish

Male Scyliorhinus canicula possess a longer and narrower mouth than females resulting in pronouced sexual dimorphism with respect to the mouth length/mouth width ratio (0.49 and 0.43, respectively). Significant sexual differences in the girth of the head and pre-oral, pre-branchial and head lengths were also recorded. Males were found to have longer teeth than females. Reasons for these differences are discussed.     

Correlated changes in the structure of the anterior pituitary gland,testes and interrenal tissue during sexual maturation of male lizards

Summary The anterior pituitary gland, testes and interrenal glands of a series of young males of the teiid Cnemidophorus l. lemniscatus (L.) have been studied by light microscopy in order to correlate the changes occurring during sexual maturation. In the testes of the smallest animals, spermatogenesis does not progress beyond primary spermatocytes and there is no differentiated interstitial tissue. In medium-sized animals, spermatids and some interstitial cells appear, and in the largest lizards, spermatogenesis is completely established and Leydig cells abound. Simultaneously with the development of the testes, interrenal glands undergo great hypertrophy and hyperplasia, especially in the peripheral reactive zone. Starting in animals of intermediate size, the anterior hypophysis exhibits a considerable hypertrophy of two rostral cell types: the chromophobic corticotrophs and the acidophilic PAS-positive cells considered to be interstitiotrophs. These cells show large, vesicular nuclei and prominent nucleoli, signs of enhanced cellular activity. The hypertrophy begins in the dorso-rostral region of the gland close to the median eminence, at the site of entry of the portal vessels. This suggests a hypothalamic influence on the function of these pars distalis cells. The scattered basophilic gonadotrophs or folliculotrophs are scarce, small, and do not vary appreciably among the animals studied. The hyperactivity of corticotrophs may account for enlargement of the interrenal glands. Testicular development is apparently related to an increased activity of interstitiotrophs but to a stable level of activity in folliculotrophs.This research forms part of project No. 31.26.S1-0244 supported by the Consejo Nacional de Investigaciones Científicas y Tecnológicas.     

Growth-related shape changes in the fetal craniofacial complex of humans (Homo sapiens) and pigtailed macaques (Macaca nemestrina): a 3D-CT comparative analysis

This study investigates whether macaques and humans possess a common pattern of relative growth during the fetal period. The fetal samples consist of 16 male pigtailed macaques (mean age, 20.5 gestational weeks) and 17 humans (9 males and 8 females; mean age, 29.5 gestational weeks). For each individual, three-dimensional coordinates of 18 landmarks on the skull were collected from three-dimensional computed tomographic (CT) reconstructed images and two-dimensional CT axial slices. Early and late groups were created from the human (early mean age, 24 weeks, N = 8; late mean age, 34 weeks, N = 9) and macaque samples (early mean age, 17.7 weeks, N = 7; late mean age, 23 weeks, N = 9). Inter- and intraspecific comparisons were made between the early and late groups. To determine if macaques and humans share a common fetal pattern of relative growth, human change in shape estimated from a comparison of early and late groups was compared to the pattern estimated between early and late macaque groups. Euclidean distance matrix analysis was used in all comparisons. Intraspecific comparisons indicate that the growing fetal skull displays the greatest amount of change along mediolateral dimensions. Changes during human growth are primarily localized to the basicranium and palate, while macaques experience localized change in the midface. Interspecific comparisons indicate that the two primate species do not share a common pattern of relative growth, and the macaque pattern is characterized by increased midfacial growth relative to humans. Our results suggest that morphological differences in the craniofacial skeleton of these species are in part established by differences in fetal growth patterns.     

Growth Rings,Phenology, Hurricane Disturbance and Climate in Cyrilla racemiflora L., a Rain Forest Tree of the Luquillo Mountains,Puerto Rico1

The growth phenology of Cyrilla racemiflora L., the dominant tree species of the montane rain forest, (subtropical lower montane rain forest, sensu Holdridge) of the Luquillo Mountains of Puerto Rico was studied intensively during 1989, and then semiannually through mid-1993 to determine the periodicity of changes in xylem structure. Four trees at 770 m were monitored for flowering, branch elongation, leaf litterfall, and xylem cell growth and differentiation in the lower stem, and these events were related to local seasonal patterns of rainfall and temperature. Hurricane Hugo defoliated study trees in September, 1989. Bud-break and branch elongation in March, 1989 were followed by earlywood xylem cell production in the lower stem in April and the onset of flowering in May. Leaf litterfall was greatest between April and June, coinciding with peak branch growth and new leaf formation. Latewood xylem was produced in December. The general phenological pattern was synchronized between trees and over study years. Vessel diameter and density were monitored along with thickness of earlywood and latewood and the former converted to vessel lumen area, a measure of xylem conductance capacity. Annual growth rings were formed with periods of earlywood and latewood production coinciding with traditional summer (rainy) and winter (dry) seasons, respectively, in the Luquillo Mountains. Hurricane defoliation was followed by heavy flowering in 1990, a year of reduced branch elongation and annual xylem ring width, and was associated with maximum vessel lumen area, as was flowering in 1989, prior to the hurricane. Hurricane Hugo provided a perturbation that, through its elicited stress response, allowed for the demonstration of the interplay between flowering, branching, structural growth of xylem, and xylem function.     

The tale of the shrinking weapon: seasonal changes in nutrition affect weapon size and sexual dimorphism,but not contemporary evolution

Sexually selected traits are often highly variable in size within populations due to their close link with the physical condition of individuals. Nutrition has a large impact on physical condition, and thus, any seasonal changes in nutritional quality are predicted to alter the average size of sexually selected traits as well as the degree of sexual dimorphism in populations. However, although traits affected by mate choice are well studied, we have a surprising lack of knowledge of how natural variation in nutrition affects the expression of sexually selected weapons and sexual dimorphism. Further, few studies explicitly test for differences in the heritability and mean‐scaled evolvability of sexually selected traits across conditions. We studied Narnia femorata (Hemiptera: Coreidae), an insect where males use their hind legs as weapons and the femurs are enlarged, to understand the extent to which weapon expression, sexual dimorphism and evolvability change across the actual range of nutrition available in the wild. We found that insects raised on a poor diet (cactus without fruit) are nearly monomorphic, whereas those raised on a high‐quality diet (cactus with ripe fruit) are distinctly sexually dimorphic via the expression of large hind leg weapons in males. Contrary to our expectations, we found little evidence of a potential for evolutionary change for any trait measured. Thus, although we show weapons are highly condition dependent, and changes in weapon expression and dimorphism could alter evolutionary dynamics, our populations are unlikely to experience further evolutionary changes under current conditions.     

Reproduction and sexual dimorphism in two populations of the polymorphic spiny lizard Sceloporus minor from Hidalgo,México

Male‐biased sexual dimorphism is extensive in New World spiny lizards (Sceloporus: Phrynosomatidae) and is particularly prominent in the polymorphic minor lizard Sceloporus minor. However, the possible relationship between patterns of sexual dimorphism and gonadal development is little known for most species. In this study, we explored aspects of sexual dimorphism in noncolor morphological traits in S. minor and characterized the gonadal cycle in males and females from each of two sites (El Enzuelado and La Manzana) in Hidalgo, México, differing in key ecological aspects linked to life‐history trait expression in other lizards. Males were generally larger than females in each population and expressed larger forms of several other morphological traits, although not all comparisons attained statistical significance. Both sexes attained reproductive maturity at a larger size in El Enzuelado relative to La Manzana, and females from El Enzuelado had larger litters. Gonadal cycles differed substantially between the two populations and suggest that reproductive activity of males and females was synchronous at El Enzuelado and asynchronous at La Manzana, an unusual pattern. Geographic variation in sex‐specific responses to environmental variables may be at least partly responsible for the exceptional species diversity of spiny lizards in México.     

浙江丽水中国石龙子的食性、两性异形和雌性繁殖

丽水分布的中国石龙子（Ｅｕｍｅｃｅｓ ｃｈｉｎｅｎｓｉｓ）摄入的食物均为无脊椎动物,分别属于环节、软体和节肢动物,涉及３０余科,成体和幼体的食物生态位宽度分别为７．２６和６．６９,成体的幼体的食物生态们重叠度为０．５９。性成熟雄性个体大于雌体。成雄和幼体的头长和头宽随体长ＳＶＬ的增长速率大于成雌,成雄头长随ＳＶＬ的增长速度显著大于幼体,成雌和幼体的头长随ＳＶＬ的增长速率无显著差异。成雄头部大于成雌