Phenotypic variability in nesting success among Osmia lignaria propinqua females in a glasshouse environment: (Hymenoptera: Megachilidae)

Abstract. 1. The phenotypic variability of several estimates of fitness among seventeen nesting female O.lignaria Cresson was examined in a glasshouse provided with abundant resources.
2. Females exhibited wide variation in: (a) number of nests and cells made and in their rate of construction, (b) the incidence of mortality of offspring, and (c) the percentage of male offspring produced (52–100%).
3. In confirmation of earlier field studies: (a) more offspring of both sexes were produced during the first half of the nesting season, (b) most female offspring were produced early in the nesting season, (c) most parental investment during the last half of the flight season was made in male offspring, (d) there was a decrease in offspring size with season, with females decreasing more than males, and (e) the ratio of female/male body weight was = 1.8. These results appear to be due to ageing rather than to a reduction in resource availability.
4. No relation between female size and fecundity was evident.
5. There was no evidence that small females tended to produce a greater proportion of the smaller sex (males) than did large females.
6. Smaller offspring of both sexes had a much greater probability of dying overwinter than did large offspring. However, where the sexes overlapped in size (large males, small females), almost all females died and almost all males survived. We hypothesize that as size of offspring produced declines with season, a greater proportion of males are produced because they have a much greater probability of surviving at small body sizes.     

Life history of Eulemur fulvus rufus from 1988-1998 in southeastern Madagascar

In this study, we compare the life-history patterns of male and female Eulemur fulvus rufus based on longitudinal data collected on individuals from two study groups from 1988-1998 in southeastern Madagascar. Mean group size was 9.5 individuals, and groups either contained more adult males than females or equal numbers of both sexes. Females reproduced for the first time between 2 and 4 years of age and reproduced each year, although the mean interbirth interval between surviving offspring was 2.1 years. An average of two adult females reproduced annually in each social group, and age and body weight may positively influence reproductive success. Females also appear to be philopatric but not female-bonded. Young natal males immigrated between 3 and 4.5 years of age and may join a new group within 612 months based on the age of emigrants. Once in a social group, they remained until old age, although a male's spatial position in the social group varied with age. Young nonnatal males were members of the social core and had the first opportunity to mate with all estrous females. Older males were peripheral to the social group and mated with females later in their cycle. We hypothesize that group size, the number of females in the group, and individual variation in reproductive success is influenced by several ecological conditions at this site: extreme variability in food availability during reproductive periods, the lack of large food patches, low plant species diversity, and small numbers of important aseasonal food sources such as Ficus species.     

Growth and mortality in juvenile European wild rabbits (Oryctolagus cuniclus)

Factors influencing the growth and mortality patterns of young were examined in a small population of European wild rabbits. Juveniles were trapped and tagged for individual identification soon after their emergence from underground nests. Age was estimated from weight on first capture and survivorship determined from the last date individuals were observed. Differences in growth rates of young born throughout the season and between the three years of the study were related to population density. Maternal social status had no significant effect on post-emergence growth rates of kittens, or on offspring survival. Young born in females' first litters of the season, in March, had significantly lower mortality than subsequent litters. Cats and stoats were regularly observed on the study site and are likely to have been major agents of juvenile mortality. There was no difference in the growth rates of the sexes, and correspondingly non-breeding adult males and females were of similar body weight. From their first days above ground juvenile males suffered significantly higher mortality than females. Differential mortality may be linked to previously reported sex differences in behaviour if the greater activity of young males made them more vulnerable to predation or attack by neighbouring territorial conspecifics.     

Body Composition Characteristics During Puberty in Girls and Boys from Eastern Austria

Puberty represents the final stage of sexual differentiation when the individual acquires reproductive capacity. Puberty is not only characterized by maturation of sexual organs and the formation of oocytes and mature spermatozoa, but also by the development of secondary sexual dimorphism. In industrialized countries, the age of puberty has decreased steadily over the last 150 years in association with improved socio-economic conditions. However, the decreased onset of puberty, especially in females, is associated with problematic changes in behaviour such as early onset of sexual activity resulting in high-risk teenage pregnancies. In our study, we analysed the association between body composition (fat tissue and fat-free body mass, estimated by BIA analyses), height, body mass index and fat distribution and signs of puberty such as the timing of menarche in 228 girls and voice breaking and facial hair growth in 191 boys ageing between 10 and 15 years. In both sexes, signs of puberty were associated, highly significantly, with body composition parameters. Nevertheless, marked differences between the two sexes were observed: Female puberty was positively associated with weight status and the absolute and relative amount of body fat, while in males, puberty was positively related with a higher amount of fat-free body mass and a decreased fat mass. Male voice breaking was significantly associated with increased stature, body weight, waist and hip circumference, lean body mass and total body water. In contrast, voice breaking was significantly negatively associated with the fat percentage, the total fat mass and the waist-to-hip ratio. Female menarche was significantly positively associated with increased body weight, weight status, waist and hip circumference and also with increased absolute and relative fat mass, relative hip circumference, lean body mass and total body water. Only the waist-to-hip ratio was significantly negatively associated with the onset of menarche.     

Differences in Patterns of Reproductive Allocation between the Sexes in Nicrophorus orbicollis

Organisms are selected to maximize lifetime reproductive success by balancing the costs of current reproduction with costs to future survival and fecundity. Males and females typically face different reproductive costs, which makes comparisons of their reproductive strategies difficult. Burying beetles provide a unique system that allows us to compare the costs of reproduction between the sexes because males and females are capable of raising offspring together or alone and carcass preparation and offspring care represent the majority of reproductive costs for both sexes. Because both sexes perform the same functions of carcass preparation and offspring care, we predict that they would experience similar costs and have similar life history patterns. In this study we assess the cost of reproduction in male Nicrophorus orbicollis and compare to patterns observed in females. We compare the reproductive strategies of single males and females that provided pre- and post-hatching parental care. There is a cost to reproduction for both males and females, but the sexes respond to these costs differently. Females match brood size with carcass size, and thus maximize the lifetime number of offspring on a given size carcass. Males cull proportionately more offspring on all carcass sizes, and thus have a lower lifetime number of offspring compared to females. Females exhibit an adaptive reproductive strategy based on resource availability, but male reproductive strategies are not adaptive in relation to resource availability.     

A genetic evaluation of mating system and determinants of individual reproductive success in Atlantic salmon (Salmo salar L.)

The primary objective of this study was to use highly polymorphic microsatellite loci to estimate individual reproductive success in Atlantic salmon based on the number of surviving juveniles (young of the year) at the population level under natural conditions. We inferred reproductive strategies adopted by both sexes by applying a maximum likelihood method to determine parent-offspring genotype relationships. A high degree of variance in individual reproductive success for both males and females was revealed. The high number of mates used by both sexes is not concordant with previous behavioral studies proposing that females are mainly monogamous in this species. We found little evidence supporting the prediction from previous reports of a positive relationship between individual size and realized reproductive success for either males or females. For both sexes, however, there was a significant correlation between the number of mates and the number of offspring. These results indicate that this species' mating system is more flexible than previously thought and suggest that factors such as potential genetic benefits or environmental uncertainty may also be driving the evolution and the plasticity of mating systems in Atlantic salmon.     

Reversed sexual dimorphism in tawny owls, Strix aluco, correlates with duty division in breeding effort

Even though most bird species with a raptorial feeding habit express varying extents of reversed sexual dimorphism (RSD: females bigger than males), the evolutionary basis for its maintenance, as well as its possible secondary consequences for the ecological adaptations of the different sexes, is debated. We studied pairs of tawny owls, Strix aluco (females 20% heavier than males), throughout the year by telemetry to test whether any inter-sexual differences in movement patterns, resource partitioning and breeding effort correlated with RSD. Females were larger than males in all body size measures and were 16% heavier than would be expected from the difference in wing length alone. In accordance with predictions from flight economics, males moved longer distances per time unit than females, in particular during the post-fledging season, when they also fed chicks more often than the females. Males had larger home ranges than females during the post-fledging period, whereas the sexes had home ranges of equal size during the non-breeding season. Until 10 days after fledging, females foraged much closer to the offspring than males, apparently balancing their distance to offspring between the needs of offspring guarding and foraging. In males, the parent–offspring distance only increased with decreasing brood condition. The sexes did not differ in habitat use or feeding habits, rendering no indications of food niche partitioning. The study provides further evidence that selection for males to be light and energetically efficient foragers is the main evolutionary force behind RSD in raptorial birds, even when the prey base is confined by territoriality.     

Socioecological aspects of human reproduction in Sub-Saharan Africa

The purpose of this study was to compare fertility outcome of two populations of northern Namibia, following different ways of subsistence. The total number of offspring, but also the number of dead and surviving offspring was compared between 236 !Kung San (91 females, 145 males) hunter gatherers and 248 Kavango (87 females, 161 males) horticultural pastoralists and a small number of Kavango people living in the urban center of Rundu. While no typical differences in fertility outcome between the study populations could be observed in males, marked differences were found for the female sample. As to be expected traditional Kavango women had given birth to a higher number of children and these children had a higher chance to survive in comparison to those of !Kung San women. On the other hand Kavango females living in urban centers reported a significantly lower number of offspring. It can be concluded that even in recent populations fertility differences according to subsistence patterns are observable.     

Mid-sagittal dimensions of cervical vertebral bodies.

A series of lateral radiographs of the cervical spinal column was evaluated in order to determine vertebral body dimensions. The sample included males (N=30) and females (N=31) 18 to 24 years old, comprising three stature percentile ranges (1-20; 40-60; 80-99) of the U.S. adult population. A two-dimensional analysis of vertebral body height (average distance between superior-inferior surgaces), depth (average distance between anteriorposterior surfaces), and area (average height X average depth) revealed minimal effects due to stature. In all subjects, average depth exceeded average height for vertebral bodies C3 through C7. Upon combining stature groups, both sexes revealed maximum average values for these dimensions at the seventh cervical vertebral body. Minimum average height occurred at C5 whereas minimum average depth was found at C3. Significant correlation (alpha greater than 0.05) was found for males between ponderal index and height and depth of the C7 vertebra. Male head weight correlated significantly with C3, C4, C5 and C6 vertebral body height and with C3, C5 and C6 vertebral body depth. For females, C7 height and C6 depth correlated significantly with ponderal index and head weight respectively. Probable biomechanical relationships of specific cervical vertebral bodies are noted     

Why make daughters larger? Maternal sex-allocation and sex-dependent selection for body size in a mass-provisioning wasp, Trypoxylon politum

Mass-provisioning wasps package maternal investment into broodcells, sealed structures that contain all the provisions necessaryfor an offspring's growth and development. Optimal sex-allocationtheory predicts that if maternal provisions determine the sizeof each offspring, and the amount of provisions available toeach offspring varies, females should allocate well-stockedbrood cells to the sex that benefits most from being large.I tested this hypothesis using observations of organ-pipe wasps,Trypoxylon politum, and dissections of their nests. A Mississippipopulation of T. politum was intensively studied from 1993 to1995. This population fit the assumptions of optimal sex-allocationmodels by Green and Brockmann and Grafen. Female weight at emergencewas 1.29 times that of males, and wing length was 1.15 timesthat of males. This discrepancy in size occurred because thevolume of parental provisions strongly influenced adult bodysize, and better-stocked brood cells were preferentially allocatedto daughters. Brood-cell volume correlated with both wing lengthand weight at emergence in both sexes, and the chance that agiven brood cell contained a female offspring increased withincreasing brood-cell volume. Fitness was positively relatedto body size for females, but I found no evidence of an advantageto large males. Although there was evidence of stabilizing selectionfor male wing length in one year, there was no evidence of anincreasing relationship between body size and fitness (directionalselection) for males in either 1993 or 1994. Female fecunditywas positively related to body size in both years, indicatingthat larger females have increased reproductive success. Therate at which females provisioned brood cells was also correlatedwith body size. Observed patterns of investment in brood cellsare quantitatively consistent with the predictions of optimalsex-allocation theory, but certain aspects of female provisioningbehavior suggest females are not following a single "optimal"strategy. Patterns of provisioning were variable among differentfemales at the study site during the same year. Large femalestended to produce larger offspring. Although Brockmann and Grafen'smodel predicts a single, population wide "switchpoint" fromthe production of male to female offspring, there was no evidencefor such a switchpoint     

Implications of individual growth status on the future sex of the European eel

Sex-related differences in growth status was demonstrated in eels Anguilla anguilla reared indoors at 17, 20 or 26° C, from the elver stage. Growth status was defined as length increase, weight increase and length–weight relationship. Eels attaining at least 10 g body weight (180–220 mm body length) were tagged with Passive Integrated Transponders (PIT). Length and weight were measured at 6-week intervals, until individuals stopped growing or had attained 150 g weight (380–450 mm). Sex-specific data from potentially undifferentiated eels were provided by retrospective classification of sex. Comparisions between sexes were made within groups graded by length or weight data from the beginning of each 6-week period. There was no consistent difference in absolute length increase between small males and females, but below 40–60 g initial body weight, males displayed on average a higher weight increase than females. Males also had lower length at weight than females, even in the smallest weight groups. Early growth status may influence the future sex of undifferentiated eels, but other approaches are needed for distinction between cause and effect.     

The relationship between cumulative number of cohabiting partners and number of children for men and women in modern Sweden

Societies with institutional monogamy may be effectively polygynous through some males having several wives in succession: serial monogamy. This study investigates the relationship between the number of cohabiting partners and the number of children in a modern society where serial monogamy is common in both sexes. Data from an investigation on cohabitation and reproduction were provided by the Swedish Statistics Bureau, where the oldest cohort were used to estimated “lifetime” number of partners and children. The analyses were repeated for the next-to-oldest cohort. About 78% males and 79% females had one partner, and about 15% of both sexes had more than one partner during their reproductive lifespan in the oldest cohort. Thus, monogamy was predominant, and serial monogamy was equally common among men and women. Serial monogamy was somewhat more frequent in the next-to-oldest cohort. Remating increased the number of offspring for males, but not for females, in both cohorts. This is in accordance with the idea of serial monogamy being a conditional reproductive strategy for males in a society with institutional monogamy. However, the reproductive costs and benefits for females of remating are obscure. In the oldest cohort of males there were significantly more unskilled laborers with no partner and no children compared to other socioeconomic categories. The same pattern, however nonsignificant, was found for the next-to-oldest cohort and, according to SSB data, is strong when even younger age groups are included. Data on socioeconomic status for females were not provided.     

Behavioral syndromes influence mating systems: floater pairs of a lizard have heavier offspring

Variation in correlated behaviors or behavioral syndromes couldhave interesting effects on mating systems, especially if thevariation in syndrome exists in both sexes. Both males and femalesof the lizard Eulamprus heatwolei display two behavioral typesof a behavioral syndrome, defined by correlations between territorial,exploratory, and predator avoidance behaviors. We tested howthis variation in behavioral syndrome influences reproductivesuccess, pairing patterns, and offspring weight. We used spatialbehavior and residency in the field to identify territorialand floater individuals. Females were relocated to the laboratoryto give birth, and all offspring, dams, and potential sireswere genotyped to determine offspring paternity. During fieldsurveys, 164 lizards were caught of which 27.5% were territorialand the rest were floaters. Paternity was assigned to 66% ofthe 104 offspring produced by 33 dams. Territorial sires fathereda greater proportion of the offspring of territorial dams thanfloater sires. Larger territorial males were more likely tosire the entire clutch or share paternity with fewer additionalsires than smaller territorial males. Floater sire size, however,did not influence the number of fathers per litter. Floaterfemales produced heavier offspring than their territorial counterparts,and offspring fathered by floaters were heavier than maternalhalf-sibs fathered by territorial males. We speculate that differencesin offspring weight may be the result of differences in yolkprovisioning by females and parent genetic compatibility.     

Do big females have big pelves?

Previous studies have shown that maternal stature is a correlate of both pelvic size and reproductive efficiency. This study addresses the issue of body size and obstetric advantage. The relationship between pelvic size and three nonpelvic measures of body size is determined for females and males. The skeletal sample consists of blacks, whites, and Native Americans. The variables include 28 measures of the pelvis, length and head diameter of the femur, and clavicular length. The coefficient of multiple determination (CMD) is computed for each pelvic measure using multiple regression, with the three nonpelvic measures serving as the independent variables. Partial correlation coefficients are also calculated between each pelvic and nonpelvic variable, while controlling for the other two nonpelvic variables. The results show that all CMDs in females and all but one CMD in males are "low," i.e., below 33%. The sexes are nonsignificantly different in their CMDs for 22 of the 28 pelvic variables; of the six variables that are significantly different, five are of the midplane. The sexes are also broadly comparable in their partial correlations. The results are explained as follows. First, the concordance between the sexes in the relationship between pelvic size and nonpelvic measures of body size is due to their genetic similarity for homologous structures. Second, as pelvic size is at the minimum at the midplane, the sexual differences in CMDs are the result of selection with respect to obstetrics. Third, four explanations for the low CMDs are discussed: 1) lack of populationally or racially specific analysis; 2) nonlinear relationship between pelvic size and nonpelvic measures of body size; 3) combination of negative allometric selection between newborn body weight and maternal stature and weight with positive selection for maternal pelvic size; and 4) hormonally induced increase in pelvic capacity during parturition.     

Age structure and body size of the Chuanxi Tree Frog Hyla annectans chuanxiensis from two different elevations in Sichuan (China)

Age, body size, and growth patterns in the subtropical anuran Hyla annectans chuanxiensis from high (Dengchigou Protection Station) and low (Lingguan Town) elevations in Baoxing County of Sichuan province (China) were described using skeletochronology. Females were significantly older than males at the low-elevation site, but there was no significant difference between the sexes at the high-elevation site. Age at sexual maturity of both males and females was 2 years at the high-elevation site, whereas males matured at 1 year and females at 2 years at the low-elevation site. Males and females from the low-elevation population reached a maximum age of 3 and 4 years, respectively, whereas males and females from the high-elevation population reached a maximum age of 4 and 5 years, respectively. At both sites, females were significantly larger than males. Females and males from the high-elevation population were larger than individuals from the low-elevation population. When the effect of age was controlled, the differences in body size of the two populations were significant only for females. Von Bertalanffy growth curves indicated that the growth rates in males was greater than in females in both populations. They also showed that the growth of both sexes slowed at an earlier age in the low-elevation population than in the high-elevation population. The findings suggest that age is a major factor underlying body size patterns for both sexes, but that the elevation of the locality affects the body size of females.     

Temporal but not spatial environmental variation drives adaptive offspring sex allocation in a plural cooperative breeder

Cooperatively breeding birds have been used frequently to study sex allocation because the adaptive value of the sexes partly depends upon the costs and benefits for parents of receiving help. I examined patterns of directional sex allocation in relation to maternal condition (Trivers-Willard hypothesis), territory quality (helper competition hypothesis), and the number of available helpers (helper repayment hypothesis) in the superb starling, Lamprotornis superbus, a plural cooperative breeder with helpers of both sexes. Superb starlings biased their offspring sex ratio in relation to prebreeding rainfall, which was correlated with maternal condition. Mothers produced relatively more female offspring in wetter years, when they were in better condition, and more male offspring in drier years, when they were in poorer condition. There was no relationship between offspring sex ratio and territory quality or the number of available helpers. Although helping was male biased, females had a greater variance in reproductive success than males. These results are consistent with the Trivers-Willard hypothesis and suggest that although females in most cooperatively breeding species make sex allocation decisions to increase their future direct reproductive success, female superb starlings appear to base this decision on their current body condition to increase their own inclusive fitness.     

Hormonal correlates of dominance in meerkats (Suricata suricatta)

In cooperatively breeding meerkats (Suricata suricatta), individuals typically live in extended family groups in which the dominant male and female are the primary reproductives, while their offspring delay dispersal, seldom breed, and contribute to the care of subsequent litters. Here we investigate hormonal differences between dominants and subordinates by comparing plasma levels of luteinizing hormone (LH), estradiol and cortisol in females, and testosterone and cortisol in males, while controlling for potential confounding factors. In both sexes, hormone levels are correlated with age. In females, levels of sex hormone also vary with body weight and access to unrelated breeding partners in the same group: subordinates in groups containing unrelated males have higher levels of LH and estradiol than those in groups containing related males only. When these effects are controlled, there are no rank-related differences in circulating levels of LH among females or testosterone among males. However, dominant females show higher levels of circulating estradiol than subordinates. Dominant males and females also have significantly higher cortisol levels than subordinates. Hence, we found no evidence that the lower levels of plasma estradiol in subordinate females were associated with high levels of glucocorticoids. These results indicate that future studies need to control for the potentially confounding effects of age, body weight, and access to unrelated breeding partners before concluding that there are fundamental physiological differences between dominant and subordinate group members.     

Growth in early life and relative body size among adult polar bears (Ursus maritimus)

Data on the body length (cm) of polar bear ( Ursus maritimus ), collected as part of a long-term ecological study in the Hudson Bay region of Canada, were examined to test the hypothesis that growth early in life influences adult body size. Data for 42 females and 30 males that were captured as two-year-olds (2.6-2.9 years), after the period of maternal care, and later recaptured as adults were analysed. On average, females increased in length by 11.2% and males by 23.1 % between captures. Relative adult body length was significantly correlated with relative length as a two-year-old in females but not in males. Thus, body length attained by two-year-olds is a weaker determinant of adult body length in males than in females. We suggest that, in comparison with females, prolonged growth beyond the period of maternal care may predispose the growth and eventual adult body size of male offspring to a greater degree of environmentally mediated variation. Furthermore, while data on maternal investment strategies in polar bears are lacking, variation in the allocation of maternal resources to cubs, within and among litters, may have a limited impact on the eventual adult body size of any surviving male offspring.     

Age structure and body size of the Chuanxi Tree Frog Hyla annectans chuanxiensis from two different elevations in Sichuan (China)

Age, body size, and growth patterns in the subtropical anuran Hyla annectans chuanxiensis from high (Dengchigou Protection Station) and low (Lingguan Town) elevations in Baoxing County of Sichuan province (China) were described using skeletochronology. Females were significantly older than males at the low-elevation site, but there was no significant difference between the sexes at the high-elevation site. Age at sexual maturity of both males and females was 2 years at the high-elevation site, whereas males matured at 1 year and females at 2 years at the low-elevation site. Males and females from the low-elevation population reached a maximum age of 3 and 4 years, respectively, whereas males and females from the high-elevation population reached a maximum age of 4 and 5 years, respectively. At both sites, females were significantly larger than males. Females and males from the high-elevation population were larger than individuals from the low-elevation population. When the effect of age was controlled, the differences in body size of the two populations were significant only for females. Von Bertalanffy growth curves indicated that the growth rates in males was greater than in females in both populations. They also showed that the growth of both sexes slowed at an earlier age in the low-elevation population than in the high-elevation population. The findings suggest that age is a major factor underlying body size patterns for both sexes, but that the elevation of the locality affects the body size of females.