Evaluating the importance of individual heterogeneity in reproduction to Weddell seal population dynamics using integral projection models.

Identifying and accounting for unobserved individual heterogeneity in vital rates in demographic models is important for estimating population-level vital rates and identifying diverse life-history strategies, but much less is known about how this individual heterogeneity influences population dynamics. We aimed to understand how the distribution of individual heterogeneity in reproductive and survival rates influenced population dynamics using vital rates from a Weddell seal population by altering the distribution of individual heterogeneity in reproduction, which also altered the distribution of individual survival rates through the incorporation of our estimate of the correlation between the two rates and assessing resulting changes in population growth. We constructed an integral projection model (IPM) structured by age and reproductive state using estimates of vital rates for a long-lived mammal that has recently been shown to exhibit large individual heterogeneity in reproduction. Using output from the IPM, we evaluated how population dynamics changed with different underlying distributions of unobserved individual heterogeneity in reproduction. Results indicate that the changes to the underlying distribution of individual heterogeneity in reproduction cause very small changes in the population growth rate and other population metrics. The largest difference in the estimated population growth rate resulting from changes to the underlying distribution of individual heterogeneity was less than 1%. Our work highlights the differing importance of individual heterogeneity at the population level compared to the individual level. Although individual heterogeneity in reproduction may result in large differences in the lifetime fitness of individuals, changing the proportion of above- or below-average breeders in the population results in much smaller differences in annual population growth rate. For a long-lived mammal with stable and high adult-survival that gives birth to a single offspring, individual heterogeneity in reproduction has a limited effect on population dynamics. We posit that the limited effect of individual heterogeneity on population dynamics may be due to canalization of life-history traits.

Individual life histories: neither slow nor fast, just diverse.

The slow-fast continuum is a commonly used framework to describe variation in life-history strategies across species. Individual life histories have also been assumed to follow a similar pattern, especially in the pace-of-life syndrome literature. However, whether a slow-fast continuum commonly explains life-history variation among individuals within a population remains unclear. Here, we formally tested for the presence of a slow-fast continuum of life histories both within populations and across species using detailed long-term individual-based demographic data for 17 bird and mammal species with markedly different life histories. We estimated adult lifespan, age at first reproduction, annual breeding frequency, and annual fecundity, and identified the main axes of life-history variation using principal component analyses. Across species, we retrieved the slow-fast continuum as the main axis of life-history variation. However, within populations, the patterns of individual life-history variation did not align with a slow-fast continuum in any species. Thus, a continuum ranking individuals from slow to fast living is unlikely to shape individual differences in life histories within populations. Rather, individual life-history variation is likely idiosyncratic across species, potentially because of processes such as stochasticity, density dependence, and individual differences in resource acquisition that affect species differently and generate non-generalizable patterns across species.

Evidence of an Absence of Inbreeding Depression in a Wild Population of Weddell Seals (Leptonychotes weddellii).

Inbreeding depression can reduce the viability of wild populations. Detecting inbreeding depression in the wild is difficult; developing accurate estimates of inbreeding can be time and labor intensive. In this study, we used a two-step modeling procedure to incorporate uncertainty inherent in estimating individual inbreeding coefficients from multilocus genotypes into estimates of inbreeding depression in a population of Weddell seals (Leptonychotes weddellii). The two-step modeling procedure presented in this paper provides a method for estimating the magnitude of a known source of error, which is assumed absent in classic regression models, and incorporating this error into inferences about inbreeding depression. The method is essentially an errors-in-variables regression with non-normal errors in both the dependent and independent variables. These models, therefore, allow for a better evaluation of the uncertainty surrounding the biological importance of inbreeding depression in non-pedigreed wild populations. For this study we genotyped 154 adult female seals from the population in Erebus Bay, Antarctica, at 29 microsatellite loci, 12 of which are novel. We used a statistical evidence approach to inference rather than hypothesis testing because the discovery of both low and high levels of inbreeding are of scientific interest. We found evidence for an absence of inbreeding depression in lifetime reproductive success, adult survival, age at maturity, and the reproductive interval of female seals in this population.

Patterns, sources, and consequences of variation in age-specific vital rates: Insights from a long-term study of Weddell seals.

Variations in the reproductive and survival abilities of individuals within a population are ubiquitous in nature, key to individual fitness, and affect population dynamics, which leads to strong interest in understanding causes and consequences of vital-rate variation. For long-lived species, long-term studies of large samples of known-age individuals are ideal for evaluating vital-rate variation. A population of Weddell seals in Erebus Bay, Antarctica, has been studied each Austral spring since the 1960s. Since 1982, all newborns have been tagged each year and multiple capture-mark-recapture (CMR) surveys have been conducted annually. Over the past 20 years, a series of analyses have built on results of earlier research by taking advantage of steady improvements in the project's long-term CMR data and available analytical methods. Here, I summarize progress made on four major topics related to variation in age-specific vital rates for females: early-life survival and age at first reproduction, costs of reproduction, demographic buffering, and demographic senescence. Multistate modelling found that age at first reproduction varies widely (4-14 years of age) and identified contrasting influences of maternal age on survival and recruitment rates of offspring. Subsequent analyses of data for females after recruitment revealed costs of reproduction to both survival and future reproduction and provided strong evidence of demographic buffering. Recent results indicated that important levels of among-individual variation exist in vital rates and revealed contrasting patterns for senescence in reproduction and survival. Sources of variation in vital rates include age, reproductive state, year, and individual. The combination of luck and individual quality results in strong variation in individual fitness outcomes: ~80% of females born in the population produce no offspring, and the remaining 20% vary strongly in lifetime reproductive output (range: 1-23 pups). Further research is needed to identify the specific environmental conditions that lead to annual variation in vital rates and to better understand the origins of individual heterogeneity. Work is also needed to better quantify the relative roles of luck, maternal effects, and environmental conditions on variation in vital rates and to learn the importance of such variation to demographic performance of offspring and on overall population dynamics.

Temporal correlations among demographic parameters are ubiquitous but highly variable across species.

Temporal correlations among demographic parameters can strongly influence population dynamics. Our empirical knowledge, however, is very limited regarding the direction and the magnitude of these correlations and how they vary among demographic parameters and species' life histories. Here, we use long-term demographic data from 15 bird and mammal species with contrasting pace of life to quantify correlation patterns among five key demographic parameters: juvenile and adult survival, reproductive probability, reproductive success and productivity. Correlations among demographic parameters were ubiquitous, more frequently positive than negative, but strongly differed across species. Correlations did not markedly change along the slow-fast continuum of life histories, suggesting that they were more strongly driven by ecological than evolutionary factors. As positive temporal demographic correlations decrease the mean of the long-run population growth rate, the common practice of ignoring temporal correlations in population models could lead to the underestimation of extinction risks in most species.

Nutrient sources for offspring formation: diet-mother and mother-offspring isotope discrimination in domesticated gallinaceous birds.

Stable isotope techniques can be used to assess nutrient acquisition and allocation strategies used to produce offspring. Before stable isotope techniques can be employed, researchers need reliable isotope discrimination values. In this context, isotope discrimination compares the difference in the isotope ratio between the maternal-offspring tissue that occurs during nutrient transfer prior to egg laying. Currently, isotope discrimination values are unknown between the maternal blood constituents - that reflect different temporal scales of integration - and downy feathers of their offspring. In this study, we experimentally derive isotope discrimination relationships between maternal diet-blood constituents for egg laying, and between maternal blood constituents-down feathers of offspring in an experiment with 3 types of domesticated gallinaceous birds raised on known diets. Our experiment is the first to report isotope discrimination values for maternal blood constituents-down of offspring in avian taxa and provides a new sampling technique that is less invasive than previously available as collecting down does not require sampling viable eggs or individuals. Future researchers can use these results to assist in identifying nutrient sources used by adult birds to produce young.

Insights from the first global population estimate of Weddell seals in Antarctica.

The Weddell seal is one of the best-studied marine mammals in the world, owing to a multidecadal demographic effort in the southernmost part of its range. Despite their occurrence around the Antarctic coastline, we know little about larger scale patterns in distribution, population size, or structure. We combined high-resolution satellite imagery from 2011, crowd-sourcing, and habitat modeling to report the first global population estimate for the species and environmental factors that influence its distribution. We estimated ~202,000 (95% confidence interval: 85,345 to 523,140) sub-adult and adult female seals, with proximate ocean depth and fast-ice variables as factors explaining spatial prevalence. Distances to penguin colonies were associated with seal presence, but only emperor penguin population size had a strong negative relationship. The small, estimated population size relative to previous estimates and the seals' nexus with trophic competitors indicates that a community ecology approach is required in efforts to monitor the Southern Ocean ecosystem.

Challenges and opportunities for comparative studies of survival rates: An example with male pinnipeds.

Survival rates are a central component of life-history strategies of large vertebrate species. However, comparative studies seldom investigate interspecific variation in survival rates with respect to other life-history traits, especially for males. The lack of such studies could be due to the challenges associated with obtaining reliable datasets, incorporating information on the 0-1 probability scale, or dealing with several types of measurement error in life-history traits, which can be a computationally intensive process that is often absent in comparative studies. We present a quantitative approach using a Bayesian phylogenetically controlled regression with the flexibility to incorporate uncertainty in estimated survival rates and quantitative life-history traits while considering genetic similarity among species and uncertainty in relatedness. As with any comparative analysis, our approach makes several assumptions regarding the generalizability and comparability of empirical data from separate studies. Our model is versatile in that it can be applied to any species group of interest and include any life-history traits as covariates. We used an unbiased simulation framework to provide "proof of concept" for our model and applied a slightly richer model to a real data example for pinnipeds. Pinnipeds are an excellent taxonomic group for comparative analysis, but survival rate data are scarce. Our work elucidates the challenges associated with addressing important questions related to broader ecological life-history patterns and how survival-reproduction trade-offs might shape evolutionary histories of extant taxa. Specifically, we underscore the importance of having high-quality estimates of age-specific survival rates and information on other life-history traits that reasonably characterize a species for accurately comparing across species.

Sources of variation in maternal allocation in a long-lived mammal.

Life history theory predicts allocation of energy to reproduction varies with maternal age, but additional maternal features may be important to the allocation of energy to reproduction. We aimed to characterize age-specific variation in maternal allocation and assess the relationship between maternal allocation and other static and dynamic maternal features. Mass measurements of 531 mothers and pups were used with Bayesian hierarchical models to explain the relationship between diverse maternal attributes and both the proportion of mass allocated by Weddell seal mothers, and the efficiency of mass transfer from mother to pup during lactation as well as the weaning mass of pups. Our results demonstrated that maternal mass was strongly and positively associated with the relative reserves allocated by a mother and a pup's weaning mass but that the efficiency of mass transfer declines with maternal parturition mass. Birthdate was positively associated with proportion mass allocation and pup weaning mass, but mass transfer efficiency was predicted to be highest at the mean birthdate. The relative allocation of maternal reserves declined with maternal age but the efficiency of mass transfer to pups increases, suggestive of selective disappearance of poor-quality mothers. These findings highlight the importance of considering multiple maternal features when assessing variation in maternal allocation.

An improved understanding of ungulate population dynamics using count data: Insights from western Montana.

Understanding the dynamics of ungulate populations is critical given their ecological and economic importance. In particular, the ability to evaluate the evidence for potential drivers of variation in population trajectories is important for informed management. However, the use of age ratio data (e.g., juveniles:adult females) as an index of variation in population dynamics is hindered by a lack of statistical power and difficult interpretation. Here, we show that the use of a population model based on count, classification and harvest data can dramatically improve the understanding of ungulate population dynamics by: 1) providing estimates of vital rates (e.g., per capita recruitment and population growth) that are easier to interpret and more useful to managers than age ratios and 2) increasing the power to assess potential sources of variation in key vital rates. We used a time series of elk (Cervus canadensis) spring count and classification data (2004 to 2016) and fall harvest data from hunting districts in western Montana to construct a population model to estimate vital rates and assess evidence for an association between a series of environmental covariates and indices of predator abundance on per capita recruitment rates of elk calves. Our results suggest that per capita recruitment rates were negatively associated with cold and wet springs, and severe winters, and positively associated with summer precipitation. In contrast, an analysis of the raw age ratio data failed to detect these relationships. Our approach based on a population model provided estimates of the region-wide mean per capita recruitment rate (mean = 0.25, 90% CI = 0.21, 0.29), temporal variation in hunting-district-specific recruitment rates (minimum = 0.09; 90% CI = [0.07, 0.11], maximum = 0.43; 90% CI = [0.38, 0.48]), and annual population growth rates (minimum = 0.83; 90% CI = [0.78, 0.87], maximum = 1.20; 90% CI = [1.11, 1.29]). We recommend using routinely collected population count and classification data and a population modeling approach rather than interpreting estimated age ratios as a substantial improvement in understanding population dynamics.

Simulation-based validation of spatial capture-recapture models: A case study using mountain lions.

Spatial capture-recapture (SCR) models have improved the ability to estimate densities of rare and elusive animals. However, SCR models have seldom been validated even as model formulations diversify and expand to incorporate new sampling methods and/or additional sources of information on model parameters. Information on the relationship between encounter probabilities, sources of additional information, and the reliability of density estimates, is rare but crucial to assessing reliability of SCR-based estimates. We used a simulation-based approach that incorporated prior empirical work to assess the accuracy and precision of density estimates from SCR models using spatially unstructured sampling. To assess the consequences of sparse data and potential sources of bias, we simulated data under six scenarios corresponding to three different levels of search effort and two levels of correlation between search effort and animal density. We then estimated density for each scenario using four models that included increasing amounts of information from harvested individuals and telemetry to evaluate the impact of additional sources of information. Model results were sensitive to the quantity of available information: density estimates based on low search effort were biased high and imprecise, whereas estimates based on high search effort were unbiased and precise. A correlation between search effort and animal density resulted in a positive bias in density estimates, though the bias decreased with increasingly informative datasets. Adding information from harvested individuals and telemetered individuals improved density estimates based on low and moderate effort but had negligible impact for datasets resulting from high effort. We demonstrated that density estimates from SCR models using spatially unstructured sampling are reliable when sufficient information is provided. Accurate density estimates can result if empirical-based simulations such as those presented here are used to develop study designs with appropriate amounts of effort and information sources.

Maladaptive nest-site selection by a sagebrush dependent species in a grazing-modified landscape.

Animals are expected to select habitats that maximize their fitness over evolutionary time scales. Yet in human-modified landscapes, habitat selection might not always lead to increased fitness because animals undervalue high-quality resources that appear less attractive than those of lower quality. In the American West, agriculture has modified landscapes, yet little is known about whether agricultural changes alter the reliability of the cues animals use to identify habitat quality; ultimately forming maladaptive breeding strategies where behavioral cues are mismatched with survival outcomes. Using the greater sage-grouse, a species highly dependent upon sagebrush landscapes, we (1) evaluated how females select nesting habitats based on sagebrush type, along with livestock grazing related linear and point features, and other biotic, abiotic characteristics, given hypothesized influences on hiding cover, microclimate and predator travel routes and perches, (2) compared habitat selection information with results for nest survival estimates to evaluate if selection appears to be adaptive or not, and (3) used our results to evaluate the most appropriate strategies for this species in a grazing-modified landscape. Nest-site selection for sagebrush type appears to be maladaptive: in the most-preferred sagebrush type, nest survival rate was one-fourth the rate realized by females nesting in the sagebrush type avoided. Nest survival was four times higher for nests placed away from (>100 m), rather than next to (1 m), the nearest fence, and survival was lower within sites with higher cow pie density (a proxy for previous grazing intensity). Live and dead grasses influenced selection and survival in opposing ways such that dead grass was selected for but resulted in reduced survival while live grass was avoided but resulted in increased survival. Results collectively provide the first empirical evidence that a specific type of sagebrush acts as an ecological trap while another sagebrush type is undervalued. These results also suggest that adding more fences to control livestock grazing systems will likely reduce sage-grouse nest survival.

Respiratory pathogens and their association with population performance in Montana and Wyoming bighorn sheep populations.

Respiratory disease caused by Mycoplasma ovipneumoniae and Pasteurellaceae poses a formidable challenge for bighorn sheep (Ovis canadensis) conservation. All-age epizootics can cause 10-90% mortality and are typically followed by multiple years of enzootic disease in lambs that hinders post-epizootic recovery of populations. The relative frequencies at which these epizootics are caused by the introduction of novel pathogens or expression of historic pathogens that have become resident in the populations is unknown. Our primary objectives were to determine how commonly the pathogens associated with respiratory disease are hosted by bighorn sheep populations and assess demographic characteristics of populations with respect to the presence of different pathogens. We sampled 22 bighorn sheep populations across Montana and Wyoming, USA for Mycoplasma ovipneumoniae and Pasteurellaceae and used data from management agencies to characterize the disease history and demographics of these populations. We tested for associations between lamb:ewe ratios and the presence of different respiratory pathogen species. All study populations hosted Pasteurellaceae and 17 (77%) hosted Mycoplasma ovipneumoniae. Average lamb:ewe ratios for individual populations where both Mycoplasma ovipneumoniae and Pasteurellaceae were detected ranged from 0.14 to 0.40. However, average lamb:ewe ratios were higher in populations where Mycoplasma ovipneumoniae was not detected (0.37, 95% CI: 0.27-0.51) than in populations where it was detected (0.25, 95% CI: 0.21-0.30). These findings suggest that respiratory pathogens are commonly hosted by bighorn sheep populations and often reduce recruitment rates; however ecological factors may interact with the pathogens to determine population-level effects. Elucidation of such factors could provide insights for management approaches that alleviate the effects of respiratory pathogens in bighorn sheep. Nevertheless, minimizing the introduction of novel pathogens from domestic sheep and goats remains imperative to bighorn sheep conservation.

Variation in the vital rates of an Antarctic marine predator: the role of individual heterogeneity.

Variation in life-history traits such as lifespan and lifetime reproductive output is thought to arise, in part, due to among-individual differences in the underlying probabilities of survival and reproduction. However, the stochastic nature of demographic processes can also generate considerable variation in fitness-related traits among otherwise-identical individuals. An improved understanding of life-history evolution and population dynamics therefore depends on evaluating the relative role of each of these processes. Here, we used a 33-yr data set with reproductive histories for 1,274 female Weddell seals from Erebus Bay, Antarctica, to assess the strength of evidence for among-individual heterogeneity in the probabilities of survival and reproduction, while accounting for multiple other sources of variation in vital rates. Our analysis used recent advances in Bayesian model selection techniques and diagnostics to directly compare model fit and predictive power between models that included individual effects on survival and reproduction to those that did not. We found strong evidence for costs of reproduction to both survival and future reproduction, with breeders having rates of survival and subsequent reproduction that were 3% and 6% lower than rates for non-breeders. We detected age-related changes in the rates of survival and reproduction, but the patterns differed for the two rates. Survival rates steadily declined from 0.92 at age 7 to 0.56 at the maximal age of 31 yr. In contrast, reproductive rates increased from 0.68 at age 7 to 0.79 at age 16 and then steadily declined to 0.37 for the oldest females. Models that included individual effects explained more variation in observed life histories and had better estimated predictive power than those that did not, indicating their importance in understanding sources of variation among individuals in life-history traits. We found that among-individual heterogeneity in survival was small relative to that for reproduction. Our study, which found patterns of variation in vital rates that are consistent with a series of predictions from life-history theory, is the first to provide a thorough assessment of variation in important vital rates for a long-lived, high-latitude marine mammal while taking full advantage of recent developments in model evaluation.

Evaluating sample size to estimate genetic management metrics in the genomics era.

Inbreeding and relationship metrics among and within populations are useful measures for genetic management of wild populations, but accuracy and precision of estimates can be influenced by the number of individual genotypes analysed. Biologists are confronted with varied advice regarding the sample size necessary for reliable estimates when using genomic tools. We developed a simulation framework to identify the optimal sample size for three widely used metrics to enable quantification of expected variance and relative bias of estimates and a comparison of results among populations. We applied this approach to analyse empirical genomic data for 30 individuals from each of four different free-ranging Rocky Mountain bighorn sheep (Ovis canadensis canadensis) populations in Montana and Wyoming, USA, through cross-species application of an Ovine array and analysis of approximately 14,000 single nucleotide polymorphisms (SNPs) after filtering. We examined intra- and interpopulation relationships using kinship and identity by state metrics, as well as FST between populations. By evaluating our simulation results, we concluded that a sample size of 25 was adequate for assessing these metrics using the Ovine array to genotype Rocky Mountain bighorn sheep herds. However, we conclude that a universal sample size rule may not be able to sufficiently address the complexities that impact genomic kinship and inbreeding estimates. Thus, we recommend that a pilot study and sample size simulation using R code we developed that includes empirical genotypes from a subset of populations of interest would be an effective approach to ensure rigour in estimating genomic kinship and population differentiation.

Assessing respiratory pathogen communities in bighorn sheep populations: Sampling realities, challenges, and improvements.

Respiratory disease has been a persistent problem for the recovery of bighorn sheep (Ovis canadensis), but has uncertain etiology. The disease has been attributed to several bacterial pathogens including Mycoplasma ovipneumoniae and Pasteurellaceae pathogens belonging to the Mannheimia, Bibersteinia, and Pasteurella genera. We estimated detection probability for these pathogens using protocols with diagnostic tests offered by a fee-for-service laboratory and not offered by a fee-for-service laboratory. We conducted 2861 diagnostic tests on swab samples collected from 476 bighorn sheep captured across Montana and Wyoming to gain inferences regarding detection probability, pathogen prevalence, and the power of different sampling methodologies to detect pathogens in bighorn sheep populations. Estimated detection probability using fee-for-service protocols was less than 0.50 for all Pasteurellaceae and 0.73 for Mycoplasma ovipneumoniae. Non-fee-for-service Pasteurellaceae protocols had higher detection probabilities, but no single protocol increased detection probability of all Pasteurellaceae pathogens to greater than 0.50. At least one protocol resulted in an estimated detection probability of 0.80 for each pathogen except Mannheimia haemolytica, for which the highest detection probability was 0.45. In general, the power to detect Pasteurellaceae pathogens at low prevalence in populations was low unless many animals were sampled or replicate samples were collected per animal. Imperfect detection also resulted in low precision when estimating prevalence for any pathogen. Low and variable detection probabilities for respiratory pathogens using live-sampling protocols may lead to inaccurate conclusions regarding pathogen community dynamics and causes of bighorn sheep respiratory disease epizootics. We recommend that agencies collect multiples samples per animal for Pasteurellaceae detection, and one sample for Mycoplasma ovipneumoniae detection from at least 30 individuals to reliably detect both Pasteurellaceae and Mycoplasma ovipneumoniae at the population-level. Availability of PCR diagnostic tests to wildlife management agencies would improve the ability to reliably detect Pasteurellaceae in bighorn sheep populations.

Coping with the loss of large, energy-dense prey: a potential bottleneck for Weddell Seals in the Ross Sea.

Extraction of Antarctic toothfish (Dissostichus mawsoni) in the Ross Sea began in 1997, following a management plan that targets the largest fish with a goal of reducing the spawning biomass by 50% over 35 yr. We investigate the potential long-term consequences of the reduced availability of this prey for Weddell seals (Leptonychotes weddellii). Energy demands in seals are acute, especially immediately following lactation, when females must recover substantial mass and cope with molting costs. We tested the hypothesis that toothfish are critically important for adult female seals during this period. Toothfish body mass is three orders of magnitude greater, and its energy density nearly double that of the most common seal prey, Antarctic silverfish (Pleuragramma antarcticum). Reduction or elimination of toothfish consumption could impair a female's ability to sufficiently recover and successfully produce a pup in the following pupping season. Our goals are to (1) illustrate mechanisms and conditions whereby toothfish depletion might plausibly affect seal population trends; (2) identify measurable parameters of the seals' ecology that may help better understand the potential negative impact of toothfish depletion on seal populations; and (3) promote a precautionary management approach for the fishery that includes monitoring of seal populations We constructed a set of inter-linked models of seal diving behavior, physiological condition, and demography based on existing information. We evaluate the effect of the following factors on seal mass recovery and intrinsic population growth rates: fishery depletion rate, daily diving limits, probability of a successful dive, and body mass recovery target. We show that loss of toothfish has the greatest potential impact on seal populations' growth rate. Under some scenarios, populations may decrease at >10% per year. Critical parameters to better understand fishery impacts include prevalence and size of toothfish in the seals' diet; the relationship between diet and the rate of mass recovery; and female breeding propensity in relation to body condition at the end of the molting period. Our results lend support to concerns about the potential negative impact of toothfish extraction in the Ross Sea; and to advocate for a precautionary management approach by the fishery.

Patterns of age-related change in reproductive effort differ in the pre-natal and post-natal periods in a long-lived mammal.

Age-related changes in maternal reproductive allocation for long-lived species are a key prediction from life-history theory. Theoretical and empirical work suggests that allocation may increase with age due to constraint (increases with experience) or restraint (increases with age in the face of declining residual reproductive value), and may decrease among the oldest aged animals due to senescence in reproductive function. Here, we use a hierarchical modelling approach to investigate the age-related patterns of change in maternal reproductive effort in the Weddell seal, a long-lived marine mammal with a protracted period of maternal care during which mothers allocate a large proportion of body mass while feeding little. We find that maternal allocation increases with age for young mothers during both the pre-natal and post-natal periods. In contrast, older mothers demonstrate a senescent decline in pre-natal allocation but allocate more of their declining resources to their offspring during the post-natal period. We also find strong evidence for the importance of individual effects in reproductive allocation among mothers: some mothers consistently produce heavier (or lighter) pups than expected. Our results indicate that maternal allocation changes over a mother's reproductive life span and that age-specific differences differ in notable ways in pre-natal and post-natal periods.

Birth dates vary with fixed and dynamic maternal features, offspring sex, and extreme climatic events in a high-latitude marine mammal.

Reproductive synchrony tends to be widespread in diverse species of plants and animals, especially at higher latitudes. However, for long-lived mammals, birth dates for different individuals can vary by weeks within a population. A mother's birth timing can reveal useful information about her reproductive abilities and have important implications for the characteristics and survival of her offspring. Despite this, our current knowledge of factors associated with variation in birth dates is modest. We used long-term data for known-age Weddell seals in Antarctica and a Bayesian hierarchical modeling approach to study how birth dates varied with fixed and temporally varying features of mothers, whether sex allocation varied with birth timing, and annual variation in birth dates. Based on birth dates for 4465 pups born to 1117 mothers aged 4-31, we found that diverse features of mothers were associated with variation in birth dates. Maternal identity was the most important among these. Unlike most studies, which have reported that birth dates occur earlier as mothers age, we found that birth dates progressively occurred earlier in the year in the early part of a mother's reproductive life, reached a minimum at age 16, and then occurred later at later ages. Birth dates were positively related to a mother's age at primiparity and recent reproductive effort. The earliest birth dates were for pups born to prime-age mothers who did not reproduce in the previous year but began reproduction early in life, suggesting that females in the best condition gave birth earlier than others. If so, our finding that male pups tended to be born earlier than females provides support for the Trivers-Willard sex-allocation model. Average birth dates were quite consistent across years, except for 2 years that had notable delays and occurred during the period when massive icebergs were present and disrupted the ecosystem.

Female Weddell seals show flexible strategies of colony attendance related to varying environmental conditions.

Many animal life cycles involve movements among different habitats to fulfill varying resource demands. There are inherent costs associated with such movements, and the decision to leave or stay at a given location ought to be motivated by the benefits associated with potential target habitats. Because movement patterns, especially those associated with reproduction, can have important implications for the success (survival, reproduction) of individual animals, and therefore a population's dynamics, it is important to identify and understand their sources of variation (environmental and individual). Here, using a mark-recapture, multistate modeling approach, we investigated a set of a priori hypotheses regarding sources and patterns of variation in breeding-colony attendance for Weddell seal (Leptonychotes weddellii) females on sabbatical from pup production. For such females, colony attendance might be motivated by predation avoidance and positive social interactions related to reproduction, but some costs, such as reduced foraging opportunities or aggressive interactions with conspecifics, might also exist. We expected these benefits and costs to vary with a female's condition and the environment. Results revealed that the probability of being absent from colonies was higher (1) in years when the extent of local sea ice was larger, (2) for the youngest and oldest individuals, and (3) for females with less reproductive experience. We also found substantial levels of residual individual heterogeneity in these rates. Based on our a priori predictions, we postulate that the decision to attend breeding colonies or not is directly influenced by an individual's physiological condition, as well as by the ice-covered distance to good foraging areas, availability of predator-free haul-out sites, and the level of negative interactions with conspecifics inside colonies. Our results support the idea that in iteroparous species, and colonial animals in particular, seasonal and temporary movements from/to reproductive sites represent flexible behavioral strategies that can play an important role in coping with environmental variability.