Sex-Biased Harvesting and Population Dynamics in Ungulates: Implications for Conservation and Sustainable Use

The consumptive use of wildlife, in particular trophy hunting and game ranching of ungulates, has been advocated as a tool for conservation in Africa. We show that these methods of harvesting differ significantly from natural predation, with trophy hunting showing extreme selection for adult males and game ranching leading to disproportionate harvests of young males. Little information, either theoretical or empirical, exists concerning the effect of these harvesting regimes on the long-term population dynamics of ungulate populations. Despite that, the potential effects of sex-skewed harvests are numerous. In this paper, we investigate one potentially deleterious effect of sex-skewed harvests. Both theory and experimental data suggest that male ungulates are limited in their absolute ability to inseminate females. Using a Leslie-Matrix model and published data on impala, we show that the interaction between sperm limitation and harvests with highly male-biased sex ratios can lead to greatly reduced female fecundity (defined as the number of young born) and population collapse. These results are robust and suggest that present methods of harvesting may not be optimal, or viable, in the long term.     

Modeling the Effects of Trophy Selection and Environmental Disturbance on a Simulated Population of African Lions

Abstract:  Tanzania is a premier destination for trophy hunting of African lions (Panthera leo) and is home to the most extensive long-term study of unhunted lions. Thus, it provides a unique opportunity to apply data from a long-term field study to a conservation dilemma: How can a trophy-hunted species whose reproductive success is closely tied to social stability be harvested sustainably? We used an individually based, spatially explicit, stochastic model, parameterized with nearly 40 years of behavioral and demographic data on lions in the Serengeti, to examine the separate effects of trophy selection and environmental disturbance on the viability of a simulated lion population in response to annual harvesting. Female population size was sensitive to the harvesting of young males (≥3 years), whereas hunting represented a relatively trivial threat to population viability when the harvest was restricted to mature males (≥6 years). Overall model performance was robust to environmental disturbance and to errors in age assessment based on nose coloration as an index used to age potential trophies. Introducing an environmental disturbance did not eliminate the capacity to maintain a viable breeding population when harvesting only older males, and initially depleted populations recovered within 15–25 years after the disturbance to levels comparable to hunted populations that did not experience a catastrophic event. These results are consistent with empirical observations of lion resilience to environmental stochasticity .     

Baboons adjust secondary sex ratio in response to predictors of sex-specific offspring survival

Summary Most theories of sex ratio adjustment assume that parents will adjust the sex ratio of births (secondary sex ratio) in a manner that maximizes offspring reproductive success (as long as this does not jeopardize parental reproductive success). Survival to maturity is typically the largest component of variance in offspring reproductive success. This should make environmental predictors of sex-specific offspring survival strong predictors of secondary sex ratio adjustment. We tested this survivorship maximization hypothesis for secondary sex ratio adjustment using data from a 17-year demographic study of 315 yellow baboon infants at Mikumi National Park, Tanzania. Sex differences were found in the degree to which several social and ecological conditions affected infant survival to 1 year. Female, but not male, infant survival was inversely correlated with birth order and the proportion of infant females in their birth cohort. Male, but not female, infant survival was inversely correlated with the amount of rainfall early in the year (January) and the proportion of infant males in their birth cohort; male survival also was positively correlated with maternal dominance rank. The consistency and timing of these effects across years suggested that such information was, in fact, available to females around the time of conception. Most importantly, social and ecological conditions that predicted improved survivorship of a given sex also were positively correlated with production of that sex. Early births were female-biased; and low January rainfall was correlated with a male-biased sex ratio, becoming increasingly female-biased as January rainfall increased. However, no sex ratio effects were correlated with maternal rank. Data supported the hypothesis that females adjusted secondary sex ratio in a manner that maximized sex-specific infant survival. This hypothesis also offered a plausible explanation for some of the contradictory data that have arisen from studies of maternal rank effects on sex ratio both within and between species.Correspondence to: S. K. Wasser     

Sex bias or equal opportunity? Patterns of maternal investment in bison

Summary In polygynous mammals, it may be adaptive for mothers to invest more in sons and/or to adjust the sex ratio of offspring in relation to body condition. Calving patterns were examined over an 8-year period (1982–1989) for a population of Bison bison in which barren females are not selectively culled. From these data, we tested predictions of the sex ratio adjustment hypothesis as well as two assumptions: (1) that offspring weight at the end of the period of parental investment (PI) is correlated with later condition, and (2) that maternal and offspring condition during the period of PI are correlated. In contrast to predictions, there was little evidence that mothers in better condition bear more sons. Short- and long-term measures of maternal condition (previous reproductive status, age, dominance status, pre-pubertal body weight, age at first reproduction, birth date, and the duration of the mother's own suckling period) were little related to offspring sex ratio, although the last calves of old females were nearly always female. Similarly, there was little evidence for sex-biased PI. Weights at about 7 months of age were greater for males than females; males also had somewhat later birth dates, suggesting either longer gestation or later conception. However, maternal reproductive costs, as measured by subsequent fecundity, weight loss, and interbirth intervals, did not vary with calf sex. Both assumptions of the model received some support. However, while maternal condition was correlated with offspring condition, there may be sex differences in investment patterns. Mothers appear better able to influence the condition of daughters than of sons. This sex difference may negate any benefit from male-biased investment.     

Population trend alters the effects of maternal dominance rank on lifetime reproductive success in yellow baboons (<Emphasis Type="Italic">Papio cynocephalus</Emphasis>)

We evaluated the association between dominance rank and lifetime reproductive success of 75 free-ranging female baboons in Mikumi National Park, Tanzania. Data were evaluated over a 22-year period that included a period of troop increase (1975–1987) associated with two troop splits in 1978 and 1979, followed by a precipitous population crash (1987–1996) where the troops successively fused back together in 1989 and 1994. Lifetime reproductive success was significantly greater for high- versus low-ranking females when examined across the entire study period. High-ranking females had a longer reproductive life span (7.4 vs 3.6 years after first birth), reached menarche earlier (4.6 vs 5.2 years), lived longer (12.0 vs 8.8 years), and had more offspring of both sexes (2.25 vs 1.33 for male offspring; 3.25 vs 0.94 for female offspring), with four times the number of offspring of each sex surviving to 4 years of age compared to low ranking females. Greater offspring production was associated with shorter interbirth intervals of dominant versus subordinate females (545 vs 723 days), partly owing to lower miscarriage rates (0.05 vs 0.2) and a shorter duration of lactation (244 vs 330 days). Rank effects were then partitioned by mothers experiencing the majority of their reproductive life prior to, versus during, the population decline. The majority of rank effects on measures of lifetime reproductive success were virtually eliminated for mothers reproducing during the troop decline, indicating that the considerable impacts of social status on lifetime reproductive success can be markedly altered by intrinsically and extrinsically mediated demographic events.Ramon Rhine is deceasedCommunicated by C. Nunn     

Selective harvesting and habitat loss produce long-term life history changes in a mouflon population.

We examined the long-term effects (28 years) of habitat loss and phenotype-based selective harvest on body mass, horn size, and horn shape of mouflon (Ovis gmelini musimon) in southern France. This population has experienced habitat deterioration (loss of 50.8% of open area) since its introduction in 1956 and unrestricted selective hunting of the largest horned males since 1973. Both processes are predicted to lead to a decrease in phenotype quality by decreasing habitat quality and by reducing the reproductive contribution of individuals carrying traits that are targeted by hunters. Body mass and body size of both sexes and horn measurements of males markedly decreased (by 3.4-38.3%) in all age classes from the 1970s. Lamb body mass varied in relation to the spatiotemporal variation of habitat closure within the hunting-free reserve, suggesting that habitat closure explains part of these changes. However, the fact that there was no significant spatial variation in body mass in the early part of the study, when a decline in phenotypic quality already had occurred, provided support for the influence of selective harvesting. We also found that the allometric relationship between horn breadth and horn length changed over the study period. For a given horn length, horn breadth was lower during the second part of the study. This result, as well as changes in horn curve diameter, supports the interpretation that selective harvesting of males based on their horn configuration had evolutionary consequences for horn shape, since this phenotypic trait is less likely to be affected by changes in habitat characteristics. Moreover, males required more time (approximately four years) to develop a desirable trophy, suggesting that trophy hunting favors the reproductive contribution of animals with slow-growing horns. Managers should exploit hunters' desire for trophy males to finance management strategies which ensure a balance between the population and its environment. However, for long-term sustainable exploitation, harvest strategy should also ensure that selectively targeted males are allowed to contribute genetically to the next generations.     

Food availability at birth limited reproductive success in historical humans

Environmental conditions in early life can profoundly affect individual development and have consequences for reproductive success. Limited food availability may be one of the reasons for this, but direct evidence linking variation in early-life nutrition to reproductive performance in adulthood in natural populations is sparse. We combined historical agricultural data with detailed demographic church records to investigate the effect of food availability around the time of birth on the reproductive success of 927 men and women born in 18th-century Finland. Our study population exhibits natural mortality and fertility rates typical of many preindustrial societies, and individuals experienced differing access to resources due to social stratification. We found that among both men and women born into landless families (i.e., with low access to resources), marital prospects, probability of reproduction, and offspring viability were all positively related to local crop yield during the birth year. Such effects were generally absent among those born into landowning families. Among landless individuals born when yields of the two main crops, rye and barley, were both below median, only 50% of adult males and 55% of adult females gained any reproductive success in their lifetime, whereas 97% and 95% of those born when both yields were above the median did so. Our results suggest that maternal investment in offspring in prenatal or early postnatal life may have profound implications for the evolutionary fitness of human offspring, particularly among those for which resources are more limiting. Our study adds support to the idea that early nutrition can limit reproductive success in natural animal populations, and provides the most direct evidence to date that this process applies to humans.     

Demographic heterogeneity, cohort selection, and population growth

Demographic heterogeneity--variation among individuals in survival and reproduction--is ubiquitous in natural populations. Structured population models address heterogeneity due to age, size, or major developmental stages. However, other important sources of demographic heterogeneity, such as genetic variation, spatial heterogeneity in the environment, maternal effects, and differential exposure to stressors, are often not easily measured and hence are modeled as stochasticity. Recent research has elucidated the role of demographic heterogeneity in changing the magnitude of demographic stochasticity in small populations. Here we demonstrate a previously unrecognized effect: heterogeneous survival in long-lived species can increase the long-term growth rate in populations of any size. We illustrate this result using simple models in which each individual's annual survival rate is independent of age but survival may differ among individuals within a cohort. Similar models, but with nonoverlapping generations, have been extensively studied by demographers, who showed that, because the more "frail" individuals are more likely to die at a young age, the average survival rate of the cohort increases with age. Within ecology and evolution, this phenomenon of "cohort selection" is increasingly appreciated as a confounding factor in studies of senescence. We show that, when placed in a population model with overlapping generations, this heterogeneity also causes the asymptotic population growth rate lambda to increase, relative to a homogeneous population with the same mean survival rate at birth. The increase occurs because, even integrating over all the cohorts in the population, the population becomes increasingly dominated by the more robust individuals. The growth rate increases monotonically with the variance in survival rates, and the effect can be substantial, easily doubling the growth rate of slow-growing populations. Correlations between parent and offspring phenotype change the magnitude of the increase in lambda, but the increase occurs even for negative parent-offspring correlations. The effect of heterogeneity in reproductive rate on lambda is quite different: growth rate increases with reproductive heterogeneity for positive parent-offspring correlation but decreases for negative parent-offspring correlation. These effects of demographic heterogeneity on lambda have important implications for population dynamics, population viability analysis, and evolution.     

Intergenerational effects of climate generate cohort variation in lizard reproductive performance

An evaluation of the link between climate and population dynamics requires understanding of climate effects both within and across generations. In ectothermic vertebrates, demographic responses to climate changes should crucially depend on balancing needs for heat and water. Here, we studied how temperature and rainfall regimes experienced before and during adulthood influenced reproductive performances (litter size, offspring size, and survival) in a natural population of the live-bearing common lizard, Lacerta vivipara, monitored continuously from 1989 to 2004. Rainfall regime, but not temperature, had both immediate and delayed effects on these reproductive performances. Rainfall during the first month of life was positively correlated with juvenile survival. Females experiencing more rainfall during gestation produced smaller neonates that showed greater survival when controlling for the positive effect of body size on survival. Furthermore, females that experienced heavier rainfall when in utero produced fewer but longer neonates during adulthood. These demographic effects of rainfall on adult reproductive traits may come from maternal effects of climate conditions and/or from delayed effects of rainfall on the environment experienced early in life. Irrespective of the precise mechanism, however, this study provides evidence of intergenerational climate effects in natural populations of an ectothermic vertebrate.     

Sexual selection and reproductive careers in mandrills (Mandrillus sphinx)

In sexually dimorphic, polygynous species, where males provide little parental care and competition between males for access to fertile females is high, sexual selection theory predicts sex differences in age-specific reproductive output and mortality profiles, and greater variance in lifetime reproductive success in males than in females. We examined age-specific reproductive output, mortality patterns and the extent and causes of variation in reproductive success for a semi-free-ranging colony of mandrills (Mandrillus sphinx, Cercopithecidae) in Franceville, Gabon, using long-term (20 year) demographic records and microsatellite parentage analysis. Although differences in the demography and feeding ecology of this closed, provisioned colony, in comparison with wild mandrills, limit interpretation of our results, sex differences in reproductive output and mortality showed the patterns predicted by sexual selection theory. Mortality was higher in males than in females after sexual maturity, and lifespan was significantly shorter in males (mean 14 year) than in females (>22 year). Age at first reproduction was significantly earlier in females (mean 4.2 year) than in males (11.6 year), and male reproductive output declined earlier. All females of breeding age produced offspring; while only 17 of 53 sexually mature males (32%) sired. Males sired a maximum of 41 offspring, versus 17 in females, and variance in male reproductive output was significantly greater than in females at all ages. The most important influence on variation in lifetime reproductive output in both sexes was joint variation between length of the breeding period and reproductive rate, due to lower reproductive rates in younger animals. Finally, social rank significantly influenced reproductive output in both sexes: high-ranking females began their reproductive careers earlier and had a higher subsequent reproductive rate than low-ranking females, while males that achieved top rank during their career sired far more offspring than males that did not.Electronic Supplementary Material Supplementary material is available for this article at     

Sex‐Biased Inbreeding Effects on Reproductive Success and Home Range Size of the Critically Endangered Black Rhinoceros

A central premise of conservation biology is that small populations suffer reduced viability through loss of genetic diversity and inbreeding. However, there is little evidence that variation in inbreeding impacts individual reproductive success within remnant populations of threatened taxa, largely due to problems associated with obtaining comprehensive pedigree information to estimate inbreeding. In the critically endangered black rhinoceros, a species that experienced severe demographic reductions, we used model selection to identify factors associated with variation in reproductive success (number of offspring). Factors examined as predictors of reproductive success were age, home range size, number of nearby mates, reserve location, and multilocus heterozygosity (a proxy for inbreeding). Multilocus heterozygosity predicted male reproductive success (p< 0.001, explained deviance >58%) and correlated with male home range size (p < 0.01, r² > 44%). Such effects were not apparent in females, where reproductive success was determined by age (p < 0.01, explained deviance 34%) as females raise calves alone and choose between, rather than compete for, mates. This first report of a 3‐way association between an individual male's heterozygosity, reproductive output, and territory size in a large vertebrate is consistent with an asymmetry in the level of intrasexual competition and highlights the relevance of sex‐biased inbreeding for the management of many conservation‐priority species. Our results contrast with the idea that wild populations of threatened taxa may possess some inherent difference from most nonthreatened populations that necessitates the use of detailed pedigrees to study inbreeding effects. Despite substantial variance in male reproductive success, the increased fitness of more heterozygous males limits the loss of heterozygosity. Understanding how individual differences in genetic diversity mediate the outcome of intrasexual competition will be essential for effective management, particularly in enclosed populations, where individuals have restricted choice about home range location and where the reproductive impact of translocated animals will depend upon the background distribution in individual heterozygosity. Efectos de la Endogamia Sesgada por el Sexo sobre el Éxito Reproductivo y el Rango del Tamaño de Hábitat del Rinoceronte Negro, Especie en Peligro Crítico     

Variations in the demographic structure and dynamics of gelada baboon populations

Summary A comparative analysis of demographic variables for three populations of gelada baboons (Theropithecus gelada) showed that these were determined by a combination of environmental, demographic and social factors interacting in complex ways. Both birth rates and survivorship were found to be adversely influenced by the severity of local climatic conditions. These independently influenced adult sex ratio, which in turn determined the proportion of multimale reproductive units in the population. Mean harem size was found to be independent of all environmental and demographic factors with the sole exception of the proportion of harem-holding males, suggesting that it is mainly a consequence of social factors related to harem fission rates. Migration rates and band fission rates were related to population growth rates, these in turn being determined largely by local mortality and birth rates.     

Mitigating the effects of climate change on the nests of sea turtles with artificial irrigation

For sea turtles, like many oviparous species, increasing temperatures during development threaten to increase embryonic mortality, alter offspring quality, and potentially create suboptimal primary sex ratios. Various methods are being implemented to mitigate the effects of climate change on reproductive success, but these methods, such as breeding programs, translocations, and shading, are often invasive and expensive. Irrigation is an alternative strategy for cooling nests that, depending on location, can be implemented relatively quickly and cheaply. However, multiple factors, including ambient conditions, nest substrate, and species characteristics, can influence irrigation success. Additionally, irrigation can vary in duration, frequency, and the volume of water applied to nests, which influences the cooling achieved and embryonic survival. Thus, it is critical to understand how to maximize cooling and manage risks before implementing irrigation as a nest-cooling strategy. We reviewed the literature on nest irrigation to examine whether artificial irrigation is feasible as a population management tool. Key factors that affected cooling were the volume of water applied and the frequency of applications. Embryonic responses varied with species, ambient conditions, and the timing of irrigation during development. Nest inundation was the key risk to a successful irrigation regime. Future irrigation regimes must identify clear targets, either primary or adult sex ratios, that maximize population viability. Monitoring population responses and adjusting the irrigation regime in response to population characteristics will be critical. Most studies reported on the manipulation of only one or two variables, further research is required to understand how altering multiple factors in an irrigation regime influences the cooling achieved and embryonic responses.     

Sex ratio and maternal rank in wild spider monkeys: when daughters disperse

Summary Data from a long-term field study of the spider monkey, Ateles paniscus, in Peru indicate that a strongly female-biased sex ratio exists from birth in this population. Of 46 infants born between July 1981 and June 1986, 12 were male, 32 were female and 2 were of undetermined sex. This effect is consistent between years as well, with more females than males born in each year of the study (Table 1). This bias is driven by the fact that low-ranking females produce daughters almost exclusively, while high-ranking females bias their investment somewhat less strongly towards sons (Table 2). The unusual pattern of female-biased maternal investment observed in this population of Ateles probably occurs for a combination of the following reasons: (1) maternal investment in individual male offspring is somewhat greater than in individual female offspring; (2) males remain with their natal groups, and the sons of high-ranking females are likely to be competitively superior to the sons of low-ranking females; (3) males compete for mates, and only the one or two most dominant males within a community are likely to achieve significant reproductive success. Two possible mechanisms of sex-ratio adjustment and the evidence for each are discussed.     

Effects of Harvesting Flowers from Shrubs on the Persistence and Abundance of Wild Shrub Populations at Multiple Spatial Extents

Abstract: Wildflower harvesting is an economically important activity of which the ecological effects are poorly understood. We assessed how harvesting of flowers affects shrub persistence and abundance at multiple spatial extents. To this end, we built a process‐based model to examine the mean persistence and abundance of wild shrubs whose flowers are subject to harvest (serotinous Proteaceae in the South African Cape Floristic Region). First, we conducted a general sensitivity analysis of how harvesting affects persistence and abundance at nested spatial extents. For most spatial extents and combinations of demographic parameters, persistence and abundance of flowering shrubs decreased abruptly once harvesting rate exceeded a certain threshold. At larger extents, metapopulations supported higher harvesting rates before their persistence and abundance decreased, but persistence and abundance also decreased more abruptly due to harvesting than at smaller extents. This threshold rate of harvest varied with species’ dispersal ability, maximum reproductive rate, adult mortality, probability of extirpation or local extinction, strength of Allee effects, and carrying capacity. Moreover, spatial extent interacted with Allee effects and probability of extirpation because both these demographic properties affected the response of local populations to harvesting more strongly than they affected the response of metapopulations. Subsequently, we simulated the effects of harvesting on three Cape Floristic Region Proteaceae species and found that these species reacted differently to harvesting, but their persistence and abundance decreased at low rates of harvest. Our estimates of harvesting rates at maximum sustainable yield differed from those of previous investigations, perhaps because researchers used different estimates of demographic parameters, models of population dynamics, and spatial extent than we did. Good demographic knowledge and careful identification of the spatial extent of interest increases confidence in assessments and monitoring of the effects of harvesting. Our general sensitivity analysis improved understanding of harvesting effects on metapopulation dynamics and allowed qualitative assessment of the probability of extirpation of poorly studied species.     

Female-biased sex allocation of offspring by an <Emphasis Type="Italic">Apodemus</Emphasis> mouse in an unstable environment

We investigated the effects of population fluctuation on the offspring’s sex allocation by a weakly polygynous mouse, Apodemus argenteus, for 3 years. In acorn-poor seasons, heavier mothers invested more in sons, and lighter mothers invested more in daughters. In acorn-rich seasons, heavier mothers invested more in daughters, and lighter mothers invested more in sons. Maternal body condition and litter size affected the sex allocation. Furthermore, there was a maternal investment trade-off between a son’s birth mass and the number of daughters. Based upon the effect of population fluctuation on the lifetime reproductive success of each sex, we proposed the new “safe bet hypothesis”. This hypothesis predicts that frequent and unpredictable change in female distribution, which is often caused by abrupt fall in food condition, favors female-biased maternal investment to offspring by polygynous mammals and is applicable to many small mammals inhabiting in unstable environments.     

Differential reproduction among female Richardson's ground squirrels and its relation to sex ratio

Summary The reproductive success of ten female Richardson's ground squirrels resident on a grassland pasture in southern Alberta in 1975 was determined by noting the number of their descendants present in 1976, 1977, and 1978.The two most successful females were the progenitors of 67% of the females resident in 1977 and of 57% of the females resident in 1978. None of the other eight females had descendants in the population in 1978. The two most successful females produced more daughters than did the other females and they did so, not by rearing larger litters, but by producing female-biased litters. The daughters of the two most successful females lived slightly longer than did the daughters of other females.Although the adult sex ratio was strongly female biased each breeding season, ranging between 0.26 and 0.42 males per female, typically all females became pregnant.Adult female offspring inherited their mother's home range and, if the mother or any female sibs were present, shared this area with them. Sons rarely remained resident in or near the natal area and adult males rarely remained resident in the same area for two consecutive years. Thus, post-weaning investments were greater in daughters than in sons.There were no conclusive correlations between sex ratio of litters and size of litters, age of the mother, previous reproductive success of the mother, population density, or the length of the overwinter period. More studies spanning several generations are required to elucidate the effects of the sex ratio of litters on the likelihood of an animal being represented by descendants in subsequent generations.     

Confronting two-sex demographic models with data

Most population dynamics models explicitly track the density of a single sex. When the operational sex ratio can vary, two-sex models may be needed to understand and predict population trajectories. Various functions have been proposed to describe the relative contributions of females and males to recruitment, and these functions can differ qualitatively in the patterns that they generate. Which mating function best describes the dynamics of real populations is not known, since alternative two-sex models have not been confronted with experimental data. We conducted the first such comparison, using laboratory populations of the bean beetle Callosobruchus maculatus. Manipulations of the operational sex ratio and total density provided strong support for a demographic model in which the birth rate was proportional to the harmonic mean of female and male densities, and females, males, and their offspring made unique contributions to density dependence. We offer guidelines for transferring this approach to other, less tractable systems in which possibilities for sex ratio manipulations are more limited. We show that informative experimental designs require strong perturbations of the operational sex ratio. The functional form of density dependence (saturating vs. over-compensatory) and the relative contributions of each sex to density dependence can both determine in which direction and at which population densities such perturbations would be most informative. Our experimental results and guidelines for design strategies promote synthesis of two-sex population dynamics theory with empirical data.     

Sigma virus and male reproductive success in Drosophila melanogaster

The risk of disease transmission can affect female mating rate, and thus sexual conflict. Furthermore, the interests of a sexually transmitted organism may align or diverge with those of either sex, potentially making the disease agent a third participant in the sexual arms race. In Drosophila melanogaster, where sexual conflict over female mating rate is well established, we investigated how a common, non-lethal virus (sigma virus) might affect this conflict. We gave uninfected females the opportunity to copulate twice in no-choice trials: either with two uninfected males, or with one male infected with sigma virus followed by an uninfected male. We assessed whether females respond behaviorally to male infection, determined whether male infection affects either female or male reproductive success, and measured offspring infection rates. Male infection status did not influence time to copulation, or time to re-mating. However, male infection did affect male reproductive success: first males sired a significantly greater proportion of offspring, as well as more total offspring, when they were infected with sigma virus. Thus viral infection may provide males an advantage in sperm competition, or, possibly, females may preferentially use infected sperm. We found no clear costs of infection in terms of offspring survival. Viral reproductive success (the number of infected offspring) was strongly correlated with male reproductive success. Further studies are needed to demonstrate whether virus-induced changes in reproductive success affect male and female lifetime fitness, and whether virus-induced changes are under male, female, or viral control.     

Sexual dimorphism, reproductive strategy, and human activities determine resource use by brown bears

Despite significant sexual dimorphism and differing reproductive strategies in carnivores, sexual segregation is rarely studied and is often overlooked in the management of wild populations. Potential nutritional constraints imposed by sexual dimorphism and differing reproductive strategies between the sexes have important implications, particularly when combined with differential effects of human activities on sex and age classes. We examined the effects of sexual dimorphism, reproductive strategies, and human activities (bear-viewing and hunting) on resource use by different sex and age classes of brown bears (Ursus arctos). Sexual segregation of habitat use and effects of experimental bear-viewing were quantified at a single site in south-central Alaska, U.S.A., by capturing, collaring, and observing brown bears at a salt marsh and salmon stream. Effects of salmon capture rate, availability of alternative salmon runs, harvest pressure, and numbers of annual visitors on sex and age class use were examined from data collected or previously published from 13 other sites. Bear-viewing sites on salmon streams where salmon capture rates were low (<4 salmon/hour) resulted in low use by adult males (<10% of all bears), except for sites with falls. However, maximum male use of viewing areas also depended on the availability of alternative salmon streams and harvest pressure. Use of habitats by females with dependent young was significantly related to the prevalence of adult males at the site. Thus, both sexual dimorphism and differing reproductive strategies led to sexual segregation in habitat use by bears. As a result of infanticide, females with young appear to prioritize avoidance of male bears over avoidance of humans when choosing habitats, in contrast to responses documented in herbivores. Because carnivores often exhibit both sexual dimorphism and infanticide, selection for sexual segregation is likely to be high. In these cases, the nutritional demands of large adult males, balanced with responses to human activity, drive dynamic temporal and spatial distributions of individuals in the population.