Spatial patterns in age- and colony-specific survival in a long-lived seabird across 14 contrasting colonies

Demographic rates such as recruitment and survival probability can vary considerably among populations of the same species due to variation in underlying environmental processes. If environmental processes are spatially correlated, nearby populations are expected to have more similar demographic rates than those further apart. Breeding populations and foraging ranges are spatially segregated in colonial seabirds, making them ideal for studying spatial patterns in demographic rates and their effects on local population dynamics. Here we explored variation in age-dependent survival probabilities across 14 colonies of Herring Gulls Larus argentatus breeding along the Dutch North Sea coast. We used long-term mark–recapture data of marked fledglings to estimate survival, and estimated spatial autocorrelation of survival probabilities. We assessed whether survival until recruitment age or until 10 years old (close to their expected lifespan) explained variation in population trajectories of each colony. Juvenile and adult survival showed a strong, but different, north-to-south gradient in survival probability, with lower juvenile but higher adult survival in northern colonies than southern colonies, whereas the spatial pattern of immature survival was less distinct. Neither recruitment nor the proportion of 10-year-old adults alive predicted whether a colony collapsed, declined, remained stable or increased. The distinct spatial pattern in survival suggests variation in regional food availability, which do not seem to drive local population dynamics. The absence of a link between survival and colony trajectories implies that connectivity between populations plays an important role affecting population dynamics.     

Estimating Survival for Elusive Juvenile Pond-Breeding Salamanders

Juvenile vital rates have important effects on population dynamics for many species, but this demographic is often difficult to locate and track. As such, we frequently lack reliable estimates of juvenile survival, which are necessary for accurately assessing population stability and potential management approaches to conserve biodiversity. We estimated survival rates for elusive juveniles of 3 species, the ringed salamander (Ambystoma annulatum), spotted salamander (A. maculatum), and small-mouthed salamander (A. texanum), using 2 approaches. First, we conducted an 11-month (2016–2017) mark-recapture study within semi-natural enclosures and used Bayesian Cormack-Jolly-Seber models to estimate survival and recapture probabilities. Second, we inferred the expected annual juvenile survival rate given published vital rates for pre-metamorphic and adult ambystomatids assuming stable population growth. For all 3 species, juvenile survival probabilities were constant across recapture occasions, whereas recapture probability estimates were time-dependent. Further, survival and recapture probabilities among study species were similar. Post-study sampling revealed that the initial study period median estimate of annual survival probability (0.39) underestimated the number of salamanders known alive at 11 months. We therefore appended approximately 1 year of opportunistic data, which produced a median annual survival probability of 0.50, encompassing salamanders that we knew to have been alive. Calculation from literature values suggested a mean annual terrestrial juvenile ambystomatid survival probability of 0.49. Similar results among our approaches indicated that juvenile survival estimates for the study species were robust and likely comparable to rates in nature. These estimates can now be confidently applied to research, monitoring, and management efforts for the study species and ecologically similar taxa. Our findings indicated that similarly robust vital rate estimates for subsets of ecologically and phylogenetically similar species can provide reasonable surrogate demographic information that can be used to reveal key factors influencing population viability for data-deficient species. © 2020 The Wildlife Society.     

Age-specific birth rates of California sea lions (Zalophus californianus) in the Gulf of California, Mexico

Estimates of demographic parameters are essential for assessing the status of populations and assigning conservation priority. In light of the difficulties associated with obtaining such estimates, vital rates are rarely available even for well-studied species. We present the first estimates of age-specific birth rates for female California sea lions ( Zalophus californianus ) >10 yr of age. These rates were estimated from the reproductive histories of five cohorts of animals branded as pups between 1980 and 1984 at Los Islotes colony in the Gulf of California, Mexico. Age-specific birth rates varied among age classes and ranged between 0.06 and 0.80. The highest birth rates were observed for females between 10 and 15 yr of age, with decreased birth rates among older females. The effect of age, year, and resighting effort were explored using logistic regression analysis. Based on Akaike Information Criteria, birth rates were best explained by female age, while year and resighting effort did not have a significant effect. The odds ratio of producing a pup decreased with age but did not change significantly for middle-aged females. Our estimates of age-specific birth rates are consistent with general patterns observed for other large vertebrates.     

Weak polygyny in California sea lions and the potential for alternative mating tactics

Female aggregation and male territoriality are considered to be hallmarks of polygynous mating systems. The development of genetic parentage assignment has called into question the accuracy of behavioral traits in predicting true mating systems. In this study we use 14 microsatellite markers to explore the mating system of one of the most behaviorally polygynous species, the California sea lion (Zalophus californianus). We sampled a total of 158 female-pup pairs and 99 territorial males across two breeding rookeries (San Jorge and Los Islotes) in the Gulf of California, Mexico. Fathers could be identified for 30% of pups sampled at San Jorge across three breeding seasons and 15% of sampled pups at Los Islotes across two breeding seasons. Analysis of paternal relatedness between the pups for which no fathers were identified (sampled over four breeding seasons at San Jorge and two at Los Islotes) revealed that few pups were likely to share a father. Thirty-one percent of the sampled males on San Jorge and 15% of the sampled males on Los Islotes were assigned at least one paternity. With one exception, no male was identified as the father of more than two pups. Furthermore, at Los Islotes rookery there were significantly fewer pups assigned paternity than expected given the pool of sampled males (p<0.0001). Overall, we found considerably lower variation in male reproductive success than expected in a species that exhibits behavior associated with strongly polygynous mating. Low variation in male reproductive success may result from heightened mobility among receptive females in the Gulf of California, which reduces the ability of males to monopolize groups of females. Our results raise important questions regarding the adaptive role of territoriality and the potential for alternative mating tactics in this species.     

Predation on an upper trophic marine predator, the Steller sea lion: evaluating high juvenile mortality in a density dependent conceptual framework

The endangered western stock of the Steller sea lion (Eumetopias jubatus)--the largest of the eared seals--has declined by 80% from population levels encountered four decades ago. Current overall trends from the Gulf of Alaska to the Aleutian Islands appear neutral with strong regional heterogeneities. A published inferential model has been used to hypothesize a continuous decline in natality and depressed juvenile survival during the height of the decline in the mid-late 1980's, followed by the recent recovery of juvenile survival to pre-decline rates. However, these hypotheses have not been tested by direct means, and causes underlying past and present population trajectories remain unresolved and controversial. We determined post-weaning juvenile survival and causes of mortality using data received post-mortem via satellite from telemetry transmitters implanted into 36 juvenile Steller sea lions from 2005 through 2011. Data show high post-weaning mortality by predation in the eastern Gulf of Alaska region. To evaluate the impact of such high levels of predation, we developed a conceptual framework to integrate density dependent with density independent effects on vital rates and population trajectories. Our data and model do not support the hypothesized recent recovery of juvenile survival rates and reduced natality. Instead, our data demonstrate continued low juvenile survival in the Prince William Sound and Kenai Fjords region of the Gulf of Alaska. Our results on contemporary predation rates combined with the density dependent conceptual framework suggest predation on juvenile sea lions as the largest impediment to recovery of the species in the eastern Gulf of Alaska region. The framework also highlights the necessity for demographic models based on age-structured census data to incorporate the differential impact of predation on multiple vital rates.     

An integrated population model sheds light on the complex population dynamics of a unique colonial breeder

Climate models forecast increasing climatic variation and more extreme events, which could increase the variability in animal demographic rates. More variable demographic rates generally lead to lower population growth and can be detrimental to wild populations, especially if the particular demographic rates affected are those to which population growth is most sensitive. We investigated the population dynamics of a metapopulation of 25 colonies of a semi-arid bird species, the sociable weaver Philetairus socius, and how it was influenced by seasonal weather during 1993–2014. We constructed an integrated population model which estimated population sizes similar to observed population counts, and allowed us to estimate annual fecundity and recruitment. Variance in fecundity contributed most to variance in population growth, which showed no trend over time. No weather variables explained overall demographic variation at the population level. However, a separate analysis of the largest colony showed a clear decline with a high extinction probability (0.05 to 0.33) within 5 years after the study period. In this colony, juvenile survival was lower when summers were hot, and adult survival was lower when winters were cold. Rainfall was also negatively correlated with adult survival. These weather effects could be due to increased physiological demands of thermoregulation and rainfall-induced breeding activity. Our results suggest that the dynamics of the population on the whole are buffered against current weather variation, as individual colonies apparently react in different ways. However, if more and increasingly extreme weather events synchronize colony dynamics, they are likely to have negative effects.     

Influence of colony size on pup fitness and survival in South American sea lions

Increased terrestrial pup mortality in small colonies due to harassment by subadult males has been proposed as a mechanism to explain the stagnation of South American sea lion populations after sealing ended. To test this hypothesis, pup survival rate was assessed in five northern Patagonia colonies with different sizes. Female diet quality as well as pup growth rate and immune status from the largest and smallest of these colonies were also assessed. Results indicated that the pup survival rate increased with colony size and pup‐to‐subadult male ratio. Furthermore, pups grew faster in the smallest colony, although female diet composition and pup immune status did not differ between the two colonies. Inverse relationship between pup growth rate and survival rate indicated that mortality was independent of food supply. In absence of terrestrial predators, infanticide by subadult males is the only mortality source other than starvation and illness and the relationship between pup survival rate and pup‐to‐subadult male ratio approached a type II functional response curve. Thus, infanticide stands as the most likely reason for the observed positive relationship between colony size and pup survival rate, supporting the hypothesis that post‐sealing population stasis was caused by inverse density dependence.     

Survival probabilities and movements of harbor seals in central California

Harbor seal numbers and population trajectories differ by location in central California. Within San Francisco Bay (SFB) counts have been relatively stable since the 1972 Marine Mammal Protection Act, but in coastal areas like Tomales Bay (TB), counts increased before stabilizing in the 1990s. Emigration, poor survival, and environmental effects have been hypothesized as contributors to differences between trajectories; however, basic demographic data were not available to evaluate these hypotheses. We monitored 32 radio‐tagged adult females (SFB n = 17, TB n = 15) for 20 mo (2011–2013), and estimated survival, resight, and movement probabilities using mark‐resight analyses and multistate mark‐resight models. Annual survival probability for both sites was 0.90 (95% CI = 0.18–0.99). Six seals were observed moving between locations resulting in an estimated probability of 0.042 (95% CI = 0.023–0.076) per month equal movement between sites. Resight probability was less in SFB relative to TB, likely due to differential haul‐out access, area surveyed, visibility, and resight effort. Because of wide confidence intervals and low precision of these first estimates of adult female harbor seal survival in California, this demographic must be further examined to dismiss its contribution to differing population trajectories. Using aerial survey data, we estimated 950 harbor seals in SFB (95% CI = 715–1,184) confirming numbers are still stable.     

Drought Impacts Survivorship and Reproductive Strategies of an Epiphyllous Leafy Liverwort in Central Amazonia

Understanding how changing precipitation patterns impact the population dynamics of Amazonian plants is necessary for predicting their long‐term survival. Most terrestrial plants are characterized by life cycles intractably slow for evaluating such demographic consequences. Due to the demands of surviving on an ephemeral substrate such as a leaf surface, epiphyllous (leaf‐inhabiting) bryophytes have some of the fastest generation times for terrestrial plants. Considering these advantages, we investigated the demographic consequences of seasonality in precipitation during one‐year of the epiphyllous bryophyte Radula flaccida Gott. (Radulaceae) in a central Amazonian rain forest. In a 1‐ha sized study area, 154 epiphyll colonies from 18 phorophytes of the understory shrub Naucleopsis ulei (Warb.) Ducke (Moraceae) were marked and colony growth, extinction, recolonization, as well as rates of sexual and asexual expression were measured in nine demographic censuses. The probability of survivorship decreased in the dry season due to the synergistic effects of both increased leaf fall (e.g., loss of occupied patches) and colony mortality on persisting leaves. Asexual expression also declined significantly during the driest months; however, sexual expression, as measured by sporophyte density, was not related to seasonality. Logistic regression probabilities also indicate a minimum threshold colony size for optimal sporophyte expression. Lower survivorship and colony growth in the dry season along with reduced sporophyte production in smaller colonies implies that both demographic stability as well as population structure of Amazonian epiphyll species may be compromised whether climate models projecting prolonged droughts for the region are accurate.     

Contrasting age-specific recruitment and survival at different spatial scales: a case study with the European storm petrel

Evolutionary studies on optimal decisions or conservation guidelines are often derived by generalising patterns from a single population, while inter‐population variability in life‐history traits is seldom considered. We investigated here how survival and recruitment probabilities changed with age at different geographical scales using the encounter histories of 5523 European storm petrels from three Mediterranean colonies, and also how our estimates of these parameters might be expected to affect population growth rates using population matrix models. We recorded similar patterns among colonies, but also important biological differences. Local survival, recruitment and breeding success increased with age at all colonies; the most distant of three colonies (Marettimo Is.) showed the largest differences. Strikingly, differences in recruitment were also found between two adjacent colonies (two caves from Benidorm Is.). Birds marked as adults from Marettimo and Benidorm colonies had a different survival, whereas we found no differences within Benidorm. Differences in survival were no longer apparent between the two islands at the end of the study following a reduction in predation by specialist gulls at Benidorm. Since birds marked as fledglings mostly recruited near the end of the study, their overall survival was high and in turn similar among colonies. Results from our population matrix models suggested that different age‐dependent patterns of demographic parameters can lead to similar population growth rates. Variability appeared to be greater for recruitment and the most sensitive parameter was adult survival. Thus conservation actions targeting this vulnerable species should focus on factors influencing adult survival. Differences in survival and recruitment among colonies could reflect the spatial heterogeneity in mortality due to predation and colony‐specific density dependent processes. Results highlight the importance of taking into account the potential spatio‐temporal heterogeneity among populations in vital rates, even in those traits that life‐history theory considers less important in driving population dynamics.     

Estimating Population Growth and Recruitment Rates Across the Range of American Common Eiders

Sound management of bird populations rests upon an adequate understanding of their population dynamics. Our study evaluated recruitment and population growth rates of 14 American common eider (Somateria mollissima dresseri) colonies from Labrador, Nova Scotia, Quebec, Canada, and Maine, USA, during various periods between 1970 and 2019. We used Pradel mark-recapture models to estimate colony-specific growth rates and the relative contributions of survival and recruitment on growth. We also validated this approach using annual nest counts (~8,000 pairs) conducted between 2003 and 2019 during down harvest operations in 3 colonies located in the Saint Lawrence estuary in Quebec. There was generally a good agreement between estimates derived using the 2 approaches. We considered that capture-recapture data were suitable to estimate population trends of common eiders in other colonies, especially for colonies where accurate nest monitoring is impaired by dense vegetation. The breeding abundance declined at major colonies in Maine and Nova Scotia and increased or was stable in Quebec and Labrador. Female survival contributed the most to population growth, but variation in recruitment among colonies was more important than variation in survival to explain population growth. Management measures should thus strive to maximize local recruitment in colonies with declining populations. The assumption that apparent survival probabilities were homogeneous throughout an individual capture history was violated at several colonies in Quebec and Labrador. Using recaptures and band recoveries, we showed that the lower apparent survival for newly marked individuals compared to females that had been recaptured at least once was caused by a difference in site fidelity rather than true survival. But <1% of recaptured females dispersed to another colony for breeding, indicating that the lower site fidelity could be related to heterogeneity in capture probability among individuals. © 2021 The Wildlife Society.     

Estimating rates of local extinction and colonization in colonial species and an extension to the metapopulation and community levels

Coloniality has mainly been studied from an evolutionary perspective, but relatively few studies have developed methods for modelling colony dynamics. Changes in number of colonies over time provide a useful tool for predicting and evaluating the responses of colonial species to management and to environmental disturbance. Probabilistic Markov process models have been recently used to estimate colony site dynamics using presence–absence data when all colonies are detected in sampling efforts. Here, we define and develop two general approaches for the modelling and analysis of colony dynamics for sampling situations in which all colonies are, and are not, detected. For both approaches, we develop a general probabilistic model for the data and then constrain model parameters based on various hypotheses about colony dynamics. We use Akaike's Information Criterion (AIC) to assess the adequacy of the constrained models. The models are parameterised with conditional probabilities of local colony site extinction and colonization. Presence–absence data arising from Pollock's robust capture–recapture design provide the basis for obtaining unbiased estimates of extinction, colonization, and detection probabilities when not all colonies are detected. This second approach should be particularly useful in situations where detection probabilities are heterogeneous among colony sites. The general methodology is illustrated using presence–absence data on two species of herons. Estimates of the extinction and colonization rates showed interspecific differences and strong temporal and spatial variations. We were also able to test specific predictions about colony dynamics based on ideas about habitat change and metapopulation dynamics. We recommend estimators based on probabilistic modelling for future work on colony dynamics. We also believe that this methodological framework has wide application to problems in animal ecology concerning metapopulation and community dynamics.     

Extreme Polygyny: Multi-seasonal “Hypergynous” Nesting in the Introduced Paper Wasp <Emphasis Type="Italic">Polistes dominulus</Emphasis>

In temperate climates, female paper wasps typically initiate new colonies in the spring. Several nest-founding tactics have been documented in Polistes species, including solitary nest initiation, joining a cooperative association, usurping an existing nest, or adopting an abandoned nest. Occasionally, exceptionally large groups of females have also been found reusing nests from the previous season. Here we report this phenomenon in introduced populations of the Eurasian species Polistes dominulus. We describe in detail the demographic and genetic characteristics of one such spring colony from Los Angeles, California, USA, which was collected with 84 associated adults and all stages of developing brood in its 613 cells. Genetic and morphological data indicate the presence of multiple reproductively active females of varying relatedness, as well as many nonbreeding females, including probable early-produced offspring. Despite some evidence of chaotic social conditions, the colony appeared to have been highly productive. Additional observations of similar colonies are needed to determine how control is maintained within such a large breeding aggregation.     

Impact of human disturbance,density, and environmental conditions on the survival probabilities of pipistrelle bat (Pipistrellus pipistrellus)

Natural and anthropogenic disturbances can strongly impact population dynamics of species and are often responsible for zoonotic emerging infectious diseases. However, long-term studies on the demographic consequences of human disturbances are unusual. We used 6 years (1995–2000) of mark-recapture data to investigate how climatic conditions, human disturbance and density affect sex- and age-specific apparent survival probabilities of the pipistrelle bat (Pipistrellus pipistrellus, Schreber 1774) in a maternity colony. Our study demonstrated that density played an important role in population dynamics of pipistrelle bat and that its effect differed with respect to age and sex. Notably, human disturbance caused a strong decline of adult female survival, suggesting that perturbations have important consequences in bat-colony dynamics. Juvenile female survival was negatively influenced by density, being considerably lower in high densities. In contrast, juvenile and adult males were apparently not affected as they had constant survival probabilities. Although climatic factors can markedly affect population dynamics of temperate insectivorous bats, in this study, the weather conditions did not influence the survival rates of pipistrelle bats. We provide the first report that demonstrates the density-dependent effect on bat survival. That is especially relevant to better understanding of the bat-population dynamics and to evaluate the consequences of human disturbance and their potential changes in the maternity colony structure.     

Contrasting Patterns of Demography and Population Viability Among Gopher Tortoise Populations in Alabama

Population viability analyses are useful tools to predict abundance and extinction risk for imperiled species. In southeastern North America, the federally threatened gopher tortoise (Gopherus polyphemus) is a keystone species in the diverse and imperiled longleaf pine (Pinus palustris) ecosystem, and researchers have suggested that tortoise populations are declining and characterized by high extinction risk. We report results from a 30-year demographic study of gopher tortoises in southern Alabama (1991–2020), where 3 populations have been stable and 3 others have declined. To better understand the demographic vital rates associated with stable and declining tortoise populations, we used a multi-state hierarchical mark-recapture model to estimate sex- and stage-specific patterns of demographic vital rates at each population. We then built a predictive population model to project population dynamics and evaluate extinction risk in a population viability context. Population structure did not change significantly in stable populations, but juveniles became less abundant in declining populations over 30 years. Apparent survival varied by age, sex, and site; adults had higher survival than juveniles, but female survival was substantially lower in declining populations than in stable ones. Using simulations, we predicted that stable populations with high female survival would persist over the next 100 years but sites with lower female survival would decline, become male-biased, and be at high risk of extirpation. Stable populations were most sensitive to changes in apparent survival of adult females. Because local populations varied greatly in vital rates, our analysis improves upon previous demographic models for northern populations of gopher tortoises by accounting for population-level variation in demographic patterns and, counter to previous model predictions, suggests that small tortoise populations can persist when habitat is managed effectively. © 2021 The Wildlife Society.     

Spatial variation in age-specific probabilities of first reproduction for Weddell seals

Spatial variation in vital rates can affect the dynamics and persistence of a population. We evaluated the prediction that age-specific probabilities of survival and first reproduction for Weddell seals would vary as a function of birth location in Erebus Bay, Antarctica. We used multi-state mark–resight models and 25 years of data to estimate demographic rates for female seals. We predicted that probabilities of survival and first reproduction would be higher for seals born at near-shore colonies or more southerly-located colonies with consistent ice conditions. Contrary to predictions, results revealed higher age-specific probabilities of first reproduction at offshore colonies relative to near-shore colonies and no spatial variation in survival rates. For 7-year old females (average age at 1st reproduction=7.6 years old) born at offshore colonies to mothers aged 10.8 years (average maternal age), probability of first reproduction was 0.43 (SE=0.07), whereas probability of first reproduction for females born at near-shore colonies was 0.30 (SE=0.05) based on estimates from our top-ranked model. Breeding probabilities following first reproduction were also higher at offshore colonies. Thus, our results (1) provide evidence of spatial variation in breeding probabilities, (2) reveal the importance of birth location on a female's vital rates, and (3) suggest that the effect persisted for many years. Birth-colony effects may be attributed to spatial variation in prey availability, or to heterogeneity in female quality in this population. If females who are superior competitors consistently chose offshore colonies for pupping, pups born at these locations may have inherited those superior qualities and displayed higher probabilities of first reproduction, relative to seals born at other colonies. Further research into physical or food-related differences among colonies may offer insight into spatial variation in breeding probabilities documented in this paper.     

Age‐related variation and temporal patterns in the survival of a long‐lived scavenger

Although senescence has been described for various fitness components in a wide range of animal species, few studies have studied senescence in long‐lived species, and little is known about its interactions with varying environmental conditions. Using a 32 year capture–mark–recapture dataset on the griffon vulture Gyps fulvus, we examined the demographic patterns of actuarial senescence and the patterns of year‐to‐year variation in survival rates. We found a significant, surprisingly late, decrease of annual survival probabilities from the age of 28 years onward and divided individual lifetimes into to three categories (juvenile, mid‐age and senescent birds). In agreement with the environmental canalization hypothesis, our analyses uncovered 1) higher temporal variation of annual survival probabilities in both the juvenile and senescent age classes compared to the mid‐age class and 2) low sensitivity of the population growth rate to the survival of both the juvenile and senescent age classes. Our results further suggested that the temporal variation in the survival of senescent birds might be related to intra‐annual changes in air temperature amplitudes. Finally, using population dynamics modeling, we revealed contrasting effects of the inclusion of the senescent age class on predicted population growth, depending on how survival rates were modeled. Altogether, our results demonstrate the existence of a class of senescent birds that exhibit distinct demographic properties compared to juvenile and mid‐age classes.     

Survival of Afro‐Palaearctic passerine migrants in western Europe and the impacts of seasonal weather variables

Populations of migratory songbirds in western Europe show considerable variation in population trends between species and regions. The demographic and environmental causes of these large‐scale patterns are poorly understood. Using data from Constant Effort mist‐netting studies, we investigated relationships between changes in abundance, adult survival and seasonal weather conditions among 35 western European populations of eight species of migratory warblers (Sylviidae). We used cross‐species and within‐species comparisons to assess whether annual variation in survival was correlated with weather conditions during passage or winter. We estimated survival using CJS mark‐recapture models accounting for variation in the proportion of transient individuals and recapture rates. Species wintering in the humid bioclimatic zone of western Africa had significantly higher annual survival probabilities than species wintering in the arid bioclimatic zone of Africa (the Sahel). Rainfall in the Sahel was positively correlated with survival in at least some populations of five species. We found substantially fewer significant relationships with indices of weather during the autumn and spring passage periods, which may be due to the use of broad‐scale indices. Annual population changes were correlated with adult survival in all of our study species, although species undergoing widespread declines showed the weakest relationships.     

Early-life density-dependence effects on growth and survival in subantarctic fur seals

Understanding the regulation of natural populations has been a long-standing research program in ecology. Current knowledge on marine mammals and seabirds is biased toward the adult component of populations and lacking are studies investigating the juvenile component. Our goal was to estimate demographic parameters on the pre-weaning stage of a subantarctic fur seal (Arctocephalus tropicalis) population on Amsterdam Island, suspected to be regulated by density-dependence. The influence of abundance on growth parameters (length and weight) and survival was assessed over a study period spanning 16 years. We evidenced a negative trend in population growth rate when density increased. Density-dependence models were favored for pup body size and mass growth. Abundance had a clear influence on body length at high population-density, pups grew slower and were smaller at weaning than pups born in years with low population density. Abundance partly explained pup body mass variation and a weak effect was detected on pre-weaning survival. The causal mechanisms may be increased competition for food resources between breeding females, leading to a reduction of maternal input to their pups. Our results suggested that pup favored survival over growth and the development of their diving abilities in order to withstand the extreme fasting periods that are characteristic of this fur seal population. This analysis provides significant insight of density-dependent processes on early-life demographic parameters of a long lived and top-predator species, and more specifically on the pre-weaning stage with important consequences for our understanding of individual long-term fitness and population dynamics.     

Incorporating movement into models of grey seal population dynamics

1. One of the most difficult problems in developing spatially explicit models of population dynamics is the validation and parameterization of the movement process. We show how movement models derived from capture-recapture analysis can be improved by incorporating them into a spatially explicit metapopulation model that is fitted to a time series of abundance data. 2. We applied multisite capture-recapture analysis techniques to photo-identification data collected from female grey seals at the four main breeding colonies in the North Sea between 1999 and 2001. The best-fitting movement models were then incorporated into state-space metapopulation models that explicitly accounted for demographic and observational stochasticity. 3. These metapopulation models were fitted to a 20-year time series of pup production data for each colony using a Bayesian approach. The best-fitting model, based on the Akaike Information Criterion (AIC), had only a single movement parameter, whose confidence interval was 82% less than that obtained from the capture-recapture study, but there was some support for a model that included an effect of distance between colonies. 4. The state-space modelling provided improved estimates of other demographic parameters. 5. The incorporation of movement, and the way in which it was modelled, affected both local and regional dynamics. These differences were most evident as colonies approached their carrying capacities, suggesting that our ability to discriminate between models should improve as the length of the grey seal time series increases.