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 .     

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.     

Factors Leading to Different Viability Predictions for a Grizzly Bear Data Set

Population viability analysis programs are being used increasingly in research and management applications, but there has not been a systematic study of the congruence of different program predictions based on a single data set. We performed such an analysis using four population viability analysis computer programs: GAPPS, INMAT, RAMAS/AGE, and VORTEX. The standardized demographic rates used in all programs were generalized from hypothetical increasing and decreasing grizzly bear ( Ursus arctos horribilis ) populations. Idiosyncracies of input format for each program led to minor differences in intrinsic growth rates that translated into striking differences in estimates of extinction rates and expected population size. In contrast, the addition of demographic stochasticity, environmental stochasticity, and inbreeding costs caused only a small divergence in viability predictions. But, the addition of density dependence caused large deviations between the programs despite our best attempts to use the same density-dependent functions. Population viability programs differ in how density dependence is incorporated, and the necessary functions are difficult to parameterize accurately. Thus, we recommend that unless data clearly suggest a particular density-dependent model, predictions based on population viability analysis should include at least one scenario without density dependence. Further, we describe output metrics that may differ between programs; development of future software could benefit from standardized input and output formats across different programs.     

Integral projection models for finite populations in a stochastic environment

Continuous types of population structure occur when continuous variables such as body size or habitat quality affect the vital parameters of individuals. These structures can give rise to complex population dynamics and interact with environmental conditions. Here we present a model for continuously structured populations with finite size, including both demographic and environmental stochasticity in the dynamics. Using recent methods developed for discrete age-structured models we derive the demographic and environmental variance of the population growth as functions of a continuous state variable. These two parameters, together with the expected population growth rate, are used to define a one-dimensional diffusion approximation of the population dynamics. Thus, a substantial reduction in complexity is achieved as the dynamics of the complex structured model can be described by only three population parameters. We provide methods for numerical calculation of the model parameters and demonstrate the accuracy of the diffusion approximation by computer simulation of specific examples. The general modeling framework makes it possible to analyze and predict future dynamics and extinction risk of populations with various types of structure, and to explore consequences of changes in demography caused by, e.g., climate change or different management decisions. Our results are especially relevant for small populations that are often of conservation concern.     

Mutation and Conservation

Mutation can critically affect the viability of small populations by causing inbreeding depression, by maintaining potentially adaptive genetic variation in quantitative characters, and through the erosion of fitness by accumulation of mildly detrimental mutations. I review and integrate recent empirical and theoretical work on spontaneous mutation and its role in population viability and conservation planning. I analyze both the maintenance of potentially adaptive genetic variation in quantitative characters and the role of detrimental mutations in increasing the extinction risk of small populations. Recent experiments indicate that the rate of production of quasineutral, potentially adaptive genetic variance in quantitative characters is an order of magnitude smaller than the total mutational variance because mutations with large phenotypic effects tend to be strongly detrimental. This implies that, to maintain normal adaptive potential in quantitative characters under a balance between mutation and random genetic drift (or among mutation, drift, and stabilizing natural selection), the effective population size should be about 5000 rather than 500 (the Franklin-Soulé number). Recent theoretical results suggest that the risk of extinction due to the fixation of mildly detrimental mutations may be comparable in importance to environmental stochasticity and could substantially decrease the long-term viability of populations with effective sizes as large as a few thousand. These findings suggest that current recovery goals for many threatened and endangered species are inadequate to ensure long-term population viability.     

Male choice,fighting ability,assortative mating and the intensity of sexual selection in the milkweed longhorn beetle,Tetraopes tetraophthalmus (Coleoptera,Cerambycidae)

Summary Over five hundred adult longhorn milkweed beetles, Tetraopes tetraophthalmus, were individually marked and their copulatory success followed for one month in a pasture of Asclepias syriaca in northern Indiana, USA. Migration of beetles from the field site was greatest from areas of low population density. Dispersal was significantly greater for males experiencing low copulatory success; a similar but nonsignificant trend was observed for females. Large males, which displayed greater site tenacity than small males, copulated more frequently than small males because of their ability to displace small males from females. Both large and small males demonstrated a preference for large females in laboratory tests. Male preference in combination with aggressive displacement of small males results in size-assortative mating which was much stronger under conditions of high population density. It contributes to variance in male reproductive success since female size is known to be correlated with fecundity and offspring viability. Variance in copulatory success is similar for males and females, suggesting that both sexes experience similar intensities of sexual selection with respect to this component of reproductive success. Futhermore, comparison of this with other studies suggests that the intensity of sexual selection among males is positively correlated with the variance in body size which appears to be under both stabilizing and directional sexual selection in males but not in females.     

A population viability analysis for the Island Fox on Santa Catalina Island,California

The island fox (Urocyon littoralis) on Santa Catalina Island is among the most imperiled species on the Channel Islands due to a recent outbreak of canine distemper virus (CDV). The western subpopulation, which was not exposed to CDV, is a crucial element in the recovery of foxes by providing a source of animals for translocation and captive breeding. Using the program VORTEX, we developed a population viability analysis for the Santa Catalina Island fox to (1) address the likelihood of population persistence, (2) estimate the current susceptibility of the population to catastrophic events, and (3) evaluate the efficacy of current restoration strategies of releasing captive bred foxes and transplanting wild animals. Overall, we found the population to be susceptible to catastrophic events; a 50% increase in mortality every 20 years was sufficient to elevate the extinction risk above 5%. Current management activities entail the transplanting of 12 juvenile foxes annually, which may reduce the viability of the western subpopulation. A minimum population size of at least 150 foxes should be maintained in each subpopulation to reduce the risk of extinction due to demographic stochasticity. Releases of translocated and captive bred animals affect the speed of recovery on the eastern half of Catalina Island, but not the probability of extinction, which is near zero under current conditions. We conducted a sensitivity analysis for demographic parameters by incrementally varying survival, fecundity and density-dependence parameters, while holding all other parameters constant. Sensitivity analyses identified mortality and mean litter size as the most sensitive parameters, while the implementation of density-dependence and environmental variation of model parameters did not seem to affect population performance. We conclude that the population of island foxes on Santa Catalina is currently at a critically low population level, but recovery of the species appears possible.     

Calculating Ne and Ne/N in age-structured populations: a hybrid Felsenstein-Hill approach

The concept of effective population size (Ne) was developed under a discrete-generation model, but most species have overlapping generations. In the early 1970s, J. Felsenstein and W. G. Hill independently developed methods for calculating Ne in age-structured populations; the two approaches produce the same answer under certain conditions and have contrasting advantages and disadvantages. Here, we describe a hybrid approach that combines useful features of both. Like Felsenstein's model, the new method is based on age-specific survival and fertility rates and therefore can be directly applied to any species for which life table data are available. Like Hill, we relax the restrictive assumption in Felsenstein's model regarding random variance in reproductive success, which allows more general application. The basic principle underlying the new method is that age structure stratifies a population into winners and losers in the game of life: individuals that live longer have more opportunities to reproduce and therefore have a higher mean lifetime reproductive success. This creates different classes of individuals within the population, and grouping individuals by age at death provides a simple means of calculating lifetime variance in reproductive success of a newborn cohort. The new method has the following features: (1) it can accommodate unequal sex ratio and sex-specific vital rates and overdispersed variance in reproductive success; (2) it can calculate effective size in species that change sex during their lifetime; (3) it can calculate Ne and the ratio Ne/N based on various ways of defining N; (4) it allows one to explore the relationship between Ne and the effective number of breeders per year (Nb), which is a quantity that genetic estimators of contemporary Ne commonly provide information about; and (5) it is implemented in freely available software (AgeNe).     

A stochastic model for estimating sustainable limits to wildlife mortality in a changing world

Human-caused mortality of wildlife is a pervasive threat to biodiversity. Assessing the population-level impact of fisheries bycatch and other human-caused mortality of wildlife has typically relied upon deterministic methods. However, population declines are often accelerated by stochastic factors that are not accounted for in such conventional methods. Building on the widely applied potential biological removal (PBR) equation, we devised a new population modeling approach for estimating sustainable limits to human-caused mortality and applied it in a case study of bottlenose dolphins affected by capture in an Australian demersal otter trawl fishery. Our approach, termed sustainable anthropogenic mortality in stochastic environments (SAMSE), incorporates environmental and demographic stochasticity, including the dependency of offspring on their mothers. The SAMSE limit is the maximum number of individuals that can be removed without causing negative stochastic population growth. We calculated a PBR of 16.2 dolphins per year based on the best abundance estimate available. In contrast, the SAMSE model indicated that only 2.3–8.0 dolphins could be removed annually without causing a population decline in a stochastic environment. These results suggest that reported bycatch rates are unsustainable in the long term, unless reproductive rates are consistently higher than average. The difference between the deterministic PBR calculation and the SAMSE limits showed that deterministic approaches may underestimate the true impact of human-caused mortality of wildlife. This highlights the importance of integrating stochasticity when evaluating the impact of bycatch or other human-caused mortality on wildlife, such as hunting, lethal control measures, and wind turbine collisions. Although population viability analysis (PVA) has been used to evaluate the impact of human-caused mortality, SAMSE represents a novel PVA framework that incorporates stochasticity for estimating acceptable levels of human-caused mortality. It offers a broadly applicable, stochastic addition to the demographic toolbox to evaluate the impact of human-caused mortality on wildlife.     

Lineage Loss in Serengeti Cheetahs: Consequences of High Reproductive Variance and Heritability of Fitness on Effective Population Size

Abstract: In natural populations, many breeders do not leave surviving offspring, and as a result many potential genetic lineages are lost. I examined lineage extinction in Serengeti cheetahs ( Acinonyx jubatus ) and found that 76% of matrilines were lost over a 25-year period. Production of future breeders was nonrandom and generally confined to a few families. Five out of 63 matrilines accounted for 45% of the total cheetah population over the course of the study. Lineage persistence is perhaps best illustrated by the variance in lifetime reproductive success ( LRS) and heritability in this parameter. In female cheetahs, variance in LRS was high, and new data show that this LRS was heritable. Variance in LRS and heritability in LRS have dramatic consequences for effective population size, N _e. I calculated N _e for cheetahs, taking into account fluctuating population size, unequal sex ratio, non-Poisson distribution of reproductive success, and heritability of fitness. The N _e was most strongly affected by variance in reproductive success and especially heritability in reproductive success. The variance N _e was 44% of the actual population size, and the inclusion of heritability further reduced N _e to only 15% of the actual population, a ratio similar to that of a social carnivore with reproductive suppression. The current cheetah population in the Serengeti is below numbers suggested by N _e estimates as sufficient to maintain sufficient genetic diversity.     

The perpetual state of emergency that sacrifices protected areas in a changing climate

A modern challenge for conservation biology is to assess the consequences of policies that adhere to assumptions of stationarity (e.g., historic norms) in an era of global environmental change. Such policies may result in unexpected and surprising levels of mitigation given future climate‐change trajectories, especially as agriculture looks to protected areas to buffer against production losses during periods of environmental extremes. We assessed the potential impact of climate‐change scenarios on the rates at which grasslands enrolled in the Conservation Reserve Program (CRP) are authorized for emergency harvesting (i.e., biomass removal) for agricultural use, which can occur when precipitation for the previous 4 months is below 40% of the normal or historical mean precipitation for that 4‐month period. We developed and analyzed scenarios under the condition that policy will continue to operate under assumptions of stationarity, thereby authorizing emergency biomass harvesting solely as a function of precipitation departure from historic norms. Model projections showed the historical likelihood of authorizing emergency biomass harvesting in any given year in the northern Great Plains was 33.28% based on long‐term weather records. Emergency biomass harvesting became the norm (>50% of years) in the scenario that reflected continued increases in emissions and a decrease in growing‐season precipitation, and areas in the Great Plains with higher historical mean annual rainfall were disproportionately affected and were subject to a greater increase in emergency biomass removal. Emergency biomass harvesting decreased only in the scenario with rapid reductions in emissions. Our scenario‐impact analysis indicated that biomass from lands enrolled in the CRP would be used primarily as a buffer for agriculture in an era of climatic change unless policy guidelines are adapted or climate‐change projections significantly depart from the current consensus.     

Incorporating landscape stochasticity into population viability analysis.

The importance of incorporating landscape dynamics into population viability analysis (PVA) has previously been acknowledged, but the need to repeat the landscape generation process to encapsulate landscape stochasticity in model outputs has largely been overlooked. Reasons for this are that (1) there is presently no means for quantifying the relative effects of landscape stochasticity and population stochasticity on model outputs, and therefore no means for determining how to allocate simulation time optimally between the two; and (2) the process of generating multiple landscapes to incorporate landscape stochasticity is tedious and user-intensive with current PVA software. Here we demonstrate that landscape stochasticity can be an important source of variance in model outputs. We solve the technical problems with incorporating landscape stochasticity by deriving a formula that gives the optimal ratio of population simulations to landscape simulations for a given model, and by providing a computer program that incorporates the formula and automates multiple landscape generation in a dynamic landscape metapopulation (DLMP) model. Using a case study of a bird population, we produce estimates of DLMP model output parameters that are up to four times more precise than those estimated from a single landscape in the same amount of total simulation time. We use the DLMP modeling software RAMAS Landscape to run the landscape and metapopulation models, though our method is general and could be applied to any PVA platform. The results of this study should motivate DLMP modelers to consider landscape stochasticity in their analyses.     

Demographic Viability of a Relict Population of the Critically Endangered Plant Borderea chouardii

Abstract:   In addition to human-caused changes in the environment, natural stochasticity may threaten species persistence, and its impact must be taken into account when priorities are established and management plans are designed. Borderea chouardii is a Tertiary relict at risk of extinction that occurs in only one location in the world, where the probability of human disturbance is low. Its persistence, therefore, is mainly linked to its response to natural threats such as stochasticity. Over 8 years I monitored up to 25% of this rupicolous small geophyte. The population had an unbalanced size structure and 90% failure in seed arrival at appropriate microhabitats, which suggests a problem with recruitment. I used matrix models to describe its population dynamics, conducted hand sowings, and performed stochastic simulations to investigate the effect of environmental stochasticity on population trend and viability. I modeled several scenarios to represent a variety of ecological situations, such as population reduction, episodic or persistent disease, and enhancement or decrease of recruitment. Population growth rate (λ) was never significantly different from unity over the study period. The risk of extinction was null over the next five centuries under current conditions. Increase of mortality and decrease of recruitment reduced stochastic population growth rate, but no factor except a persistent increase of 10% mortality resulted in extinction. These results are the consequence of the plant's extremely long life span (over 300 years) and low temporal variability of key vital rates. Even though hand sowing significantly increased the stochastic population growth rate, other approaches may be more important for the persistence of this species. The extremely slow capacity for recovery following disturbances renders habitat preservation essential. In addition, the founding of new populations would reduce the risk associated with habitat destruction.     

Population Constraints Associated with the Use of Black Rhinos as an Umbrella Species for Desert Herbivores

The numerous tactics used to conserve biodiversity include the designation of protected areas, political change, and research and education, the latter involving paradigms such as insular biogeography and the "umbrella species concept." In Namibia lands removed from national park status in 1970 and currently under the jurisdiction of indigenous people now contain one of the few unfenced populations of black rhinos (Diceros bicornis) remaining in Africa. Theory predicts that the protection of umbrella species will ensure the survival of other biota that require(s) less space. To gauge how well biodiversity might be retained by examining the spatial needs of a small population of black rhinos, I used data gathered under various ecological conditions to estimate mean and minimum population sizes of six large herbivores of the Namib Desert ranging in size from giraffe (Giraffa camelopardalis) to springbok (Antidorcas marsupialis) and ostrich (Struthio camelus). My results indicate that annual differences in rainfall, both within and between seasons, resulted in wide fluctuations in herbivore population sizes for all species except rhino. Although other herbivores switched to areas of higher rainfall, rhinos did not. The data suggest that under conditions of extreme environmental variance the space used by rhinos alone was unlikely to assure the existence of populations of other species in excess of 250 individuals. Fifty percent of the species failed to exceed 150 individuals 50% of the time and one third of the species never attained populations in excess of 50 individuals. However, by employing assumptions about the spatial needs of rhino populations numbering up to 100 individuals, the mean minimum population sizes attained by any of four desert herbivores is 535. A future challenge in using rhinos and other large-bodied species as umbrellas for organisms of either similar or dissimilar trophic levels will be the refinement of estimates of population viability.     

A Population Viability Analysis for African Elephant (Loxodonta africana): How Big Should Reserves Be?

We present an age-structured, density-dependent model of elephant population dynamics in a fluctuating environment, drawing primarily upon the life history parameters obtained from studies in semi-arid land at Tsavo National Park, Kenya. Density regulation occurs by changes in the age of first reproduction and calving interval. We model environmental stochasticity with drought events affecting sex- and age-specific survivorships. Results indicate a maximum population growth rate of 3% per year and an equilibrium elephant density of 3.1/mile². Analysis of the demographic results and their sensitivity to changes in juvenile survivorship and drought frequencies, supported by genetic considerations, suggests that in semi-arid regions a minimum reserve size of 1000 mile² is necessary to attain a 99% probability of population persistence for 1000 years. The effect of age-independent culling on population viability is also analyzed.     

Determinants of reproductive success of the White-browed Sparrow Weaver,Plocepasser mahali

Summary Variation in reproductive success among 26 communal groups in a sampled population of Plocepasser mahali (White-browed Sparrow Weaver) was studied over a 3-year period in Zambia, Africa. Potential determinants of reproductive success, namely resource variables and group size, were examined and statistically analyzed for their significance in explaining annual variance in reproductive success among these groups. Resource variables included abundance of grass seeds (dry season food) and grasshoppers (wet season food), nest tree quality, and percentage availability of preferred feeding cover. Only the latter two proved appropriate for this analysis. Patterns of utilization did not correlate positively with food abundance, and grasshopper abundance fluctuated too much among the groups in a given year to be treated as a stable variable.From an analysis of multiple correlation coefficients in a stepwise multiple regression model, both group size and ground cover explained independently of each other 20%–30% of the variance in annual production of young surviving 6 months. Explained variance by these two variables also revealed that their relative importance varied considerably between years. Hypotheses are offered to explain the possible causal mechanisms these variables may have in influencing intergroup reproductive success and the possible reasons why vagaries in explained variance were observed. It is suggested that effects of group size and habitat quality may be more important than age-specific effects in modeling population growth and regulation for species like P. mahali.     

Mast seeding under increasing drought: results from a long-term data set and from a rainfall exclusion experiment

Mast seeding, the synchronous, highly variable seed production among years, is very common in tree species, but there is no consensus about its main causes and the main environmental factors affecting it. In this study, we first analyze a long-term data set on reproductive and vegetative growth of Quercus ilex in a mediterranean woodland in order to identify the main environmental drivers of interannual variation in flower and seed production and contrast the impact of climate vs. adaptive factors as main causes of masting. Second, we conducted an experiment of rainfall exclusion to evaluate the effects of an increasing drought (simulating predictions of global change models) on both reproductive processes. The annual seed crop was always affected by environmental factors related to the precipitation pattern, these abiotic factors disrupting the fruiting process at different periods of time. Seed production was strongly dependent upon water availability for the plant at initial (spring) and advanced (summer) stages of the acorn maturation cycle, whereas the final step of seed development was negatively affected by the frequency of torrential-rain events. We also found clear evidence that seed masting in the study species is not only regulated by selective endogenous rhythms, but is mainly a physiological response to the variable environment. Our results from the rainfall exclusion experiment corroborated the conclusions obtained from the 26-year fruiting record and demonstrated that the high interannual variation in seed crop was mainly determined by the success in seed development rather than by the flowering effort. Under a global change scenario, it could be expected that the drier conditions predicted by climate models reinforce the negative effects of summer drought on seed production, leading to negative consequences for tree recruitment and forest dynamics.     

Effects of Social Structure and Prey Dynamics on Extinction Risk in Gray Wolves

Extinction models based on diffusion theory generally fail to incorporate two important aspects of population biology—social structure and prey dynamics. We include these aspects in an individual-based extinction model for small, isolated populations of the gray wolf (Canis lupus). Our model predicts mean times to extinction significantly longer than those predicted by more general (diffusion) models. According to our model, an isolated population of 50 wolves has a 95% chance of surviving just 9 years and only a 30% chance of surviving beyond 100 years. Reflecting the influence of social structure, a wolf population initially comprising 50 individuals is expected to persist only a few years longer, on average (71 years), than is a population initially comprising just a single reproductive pair (62 years). In contrast, substantially greater average prey abundance leads to dramatically longer expected persistence times. Autocorrelated prey dynamics result in a more complex distribution of extinction times than predicted by many extinction models. We contend that demographic stochasticity may pose the greatest threat to small, isolated wolf populations, although environmental stochasticity and genetic effects may compound this threat. Our work highlights the importance of considering social structure and resource dynamics in the development of population viability analyses.     

Male-biased sex ratios in New Zealand fur seal pups relative to environmental variation

A number of models have been proposed to provide adaptive explanations of sex-ratio variation in mammals. Two models have been applied commonly to primates and ungulates with varying success—the Trivers-Willard (TW) hypothesis, and the local resource competition (LRC) hypothesis. For polygynous, sexually dimorphic mammals, where males are larger and disperse more readily, these models predict opposite outcomes of sex-ratio adjustment within the same environmental context (high-resource years: TW—more sons; LRC—more daughters). However, many of the predictions of these two models can vary depending on factors influencing resource availability, such as environmental stochasticity, resource predictability, and population density. The New Zealand fur seal (Arctocephalus forsteri) is a polygynous mammal showing marked sexual dimorphism (larger males), with higher variation in male reproductive success expected. We provide clear evidence of male-biased sex ratios from a large sample of A. forsteri pups captured around South Island, New Zealand during 1996/1998, even after accounting for a sex bias in capture probability. The extent of the bias depended upon year and, in 1998, strong climatic perturbations (El Niño/Southern Oscillation, ENSO) probably reduced food availability. Significant male-biased sex ratios were found in all years; however, there was a significant decline in the male bias in 1998. There was no relationship between sex ratio and population density. We suggest that the sex-ratio bias resulted from the production of relatively more male pups. Under the density-independent scenario, the strong male bias in A. forsteri sex ratios is support for the TW model within an environment of high resource predictability. We suggest that some plasticity in the determination of pup sex among years is a mechanism by which A. forsteri females in New Zealand, and perhaps other otariid seals, can maximise fitness benefits when living in regions of high, yet apparently predictable, environmental variability. We also suggest that much of the inconsistency in the reported sex ratios for otariid seals results from the complex interaction of population density and environmental stochasticity influencing relative food availability over time.     

Predicting the Deleterious Effects of Mutation Load in Fragmented Populations

Abstract:  Human-induced habitat fragmentation constitutes a major threat to biodiversity. Both genetic and demographic factors combine to drive small and isolated populations into extinction vortices. Nevertheless, the deleterious effects of inbreeding and drift load may depend on population structure, migration patterns, and mating systems and are difficult to predict in the absence of crossing experiments. We performed stochastic individual-based simulations aimed at predicting the effects of deleterious mutations on population fitness (offspring viability and median time to extinction) under a variety of settings (landscape configurations, migration models, and mating systems) on the basis of easy-to-collect demographic and genetic information. Pooling all simulations, a large part (70%) of variance in offspring viability was explained by a combination of genetic structure ( F_ST ) and within-deme heterozygosity ( H_S ). A similar part of variance in median time to extinction was explained by a combination of local population size ( N ) and heterozygosity ( H_S ). In both cases the predictive power increased above 80% when information on mating systems was available. These results provide robust predictive models to evaluate the viability prospects of fragmented populations.