Multievent Capture-recapture Models Research Articles

Multi-event capture-recapture models estimate the diagnostic performance of serological tests for myxoma and rabbit haemorrhagic disease viruses in the absence of reference samples.

Estimation of the diagnostic performance of serological tests often relies on another test assumed as a reference or on samples of known infection status, yet both are seldom available for emerging pathogens in wildlife. Longitudinal disease serological data can be analysed through multi-event capture-mark-recapture (MECMR) models accounting for the uncertainty in state assignment, allowing us to estimate epidemiological parameters such as incidence and mortality. We hypothesized that by estimating the uncertainty in state assignment, MECMR models estimate the diagnostic performance of serological tests for rabbit haemorrhagic disease virus (RHDV) and myxoma virus (MYXV). We evaluated this hypothesis on longitudinal serological data of three tests of RHDV and one test of MYXV in two populations of the European rabbit (Oryctolagus cuniculus algirus). First, we selected the optimal cut-off threshold for each test using finite mixture models, a reference method not relying on reference tests or samples. Second, we used MECMR models to compare the diagnostic sensitivity (Se) and specificity (Sp) of the three tests for RHDV. Third, we compared the estimates of diagnostic performance by MECMR and finite mixture models across a range of cut-off values. The MECMR models showed that the RHDV test employing GI.2 antigens (Se: 100%) outperformed two tests employing GI.1 antigens (Se: 21.7% ± 8.6% and 8.7% ± 5.9%). At their selected cut-offs (2.0 for RHDV GI.2 and 2.4 for MYXV), the estimates of Se and Sp were concordant between the MECMR and finite mixture models. Over the duration of the study (May 2018 to September 2020), the monthly survival of European rabbits seropositive for MYXV was significantly higher than that of seronegative rabbits (82.7% ± 4.9%versus 61.5% ± 12.7%) at the non-fenced site. We conclude that MECMR models can reliably estimate the diagnostic performance of serological tests for RHDV and MYXV in European rabbits. This conclusion could extend to other diagnostic tests and host-pathogen systems. Longitudinal disease surveillance data analysed through MECMR models allow the validation of diagnostic tests for emerging pathogens in novel host species while simultaneously estimating epidemiological parameters.

Episodes of opposing survival and reproductive selection cause strong fluctuating selection on seasonal migration versus residence.

Quantifying temporal variation in sex-specific selection on key ecologically relevant traits, and quantifying how such variation arises through synergistic or opposing components of survival and reproductive selection, is central to understanding eco-evolutionary dynamics, but rarely achieved. Seasonal migration versus residence is one key trait that directly shapes spatio-seasonal population dynamics in spatially and temporally varying environments, but temporal dynamics of sex-specific selection have not been fully quantified. We fitted multi-event capture-recapture models to year-round ring resightings and breeding success data from partially migratory European shags (Phalacrocorax aristotelis) to quantify temporal variation in annual sex-specific selection on seasonal migration versus residence arising through adult survival, reproduction and the combination of both (i.e. annual fitness). We demonstrate episodes of strong and strongly fluctuating selection through annual fitness that were broadly synchronized across females and males. These overall fluctuations arose because strong reproductive selection against migration in several years contrasted with strong survival selection against residence in years with extreme climatic events. These results indicate how substantial phenotypic and genetic variation in migration versus residence could be maintained, and highlight that biologically important fluctuations in selection may not be detected unless both survival selection and reproductive selection are appropriately quantified and combined.

Differential survival throughout the full annual cycle of a migratory bird presents a life-history trade-off.

Long-distance migrations are among the most physically demanding feats animals perform. Understanding the potential costs and benefits of such behaviour is a fundamental question in ecology and evolution. A hypothetical cost of migration should be outweighed by higher productivity and/or higher annual survival, but few studies on migratory species have been able to directly quantify patterns of survival throughout the full annual cycle and across the majority of a species' range. Here, we use telemetry data from 220 migratory Egyptian vultures Neophron percnopterus, tracked for 3,186 bird months and across approximately 70% of the species' global distribution, to test for differences in survival throughout the annual cycle. We estimated monthly survival probability relative to migration and latitude using a multi-event capture-recapture model in a Bayesian framework that accounted for age, origin, subpopulation and the uncertainty of classifying fates from tracking data. We found lower survival during migration compared to stationary periods (β=-0.816; 95% credible interval: -1.290 to -0.318) and higher survival on non-breeding grounds at southern latitudes (<25°N; β=0.664; 0.076-1.319) compared to on breeding grounds. Survival was also higher for individuals originating from Western Europe (β=0.664; 0.110-1.330) as compared to further east in Europe and Asia, and improved with age (β=0.030; 0.020-0.042). Anthropogenic mortalities accounted for half of the mortalities with a known cause and occurred mainly in northern latitudes. Many juveniles drowned in the Mediterranean Sea on their first autumn migration while there were few confirmed mortalities in the Sahara Desert, indicating that migration barriers are likely species-specific. Our study advances the understanding of important fitness trade-offs associated with long-distance migration. We conclude that there is lower survival associated with migration, but that this may be offset by higher non-breeding survival at lower latitudes. We found more human-caused mortality farther north, and suggest that increasing anthropogenic mortality could disrupt the delicate migration trade-off balance. Research to investigate further potential benefits of migration (e.g. differential productivity across latitudes) could clarify how migration evolved and how migrants may persist in a rapidly changing world.

Non-Invasive Molecular Survey of Sarcoptic Mange in Wildlife: Diagnostic Performance in Wolf Faecal Samples Evaluated by Multi-Event Capture-Recapture Models.

Sarcoptic mange is globally enzootic, and non-invasive methods with high diagnostic specificity for its surveillance in wildlife are lacking. We describe the molecular detection of Sarcoptes scabiei in non-invasively collected faecal samples, targeting the 16S rDNA gene. We applied this method to 843 Iberian wolf Canis lupus signatus faecal samples collected in north-western Portugal (2006–2018). We further integrated this with serological data (61 samples from wolf and 20 from red fox Vulpes vulpes, 1997–2019) in multi-event capture–recapture models. The mean predicted prevalence by the molecular analysis of wolf faecal samples from 2006–2018 was 7.2% (CI95 5.0–9.4%; range: 2.6–11.7%), highest in 2009. The mean predicted seroprevalence in wolves was 24.5% (CI95 18.5–30.6%; range: 13.0–55.0%), peaking in 2006–2009. Multi-event capture–recapture models estimated 100% diagnostic specificity and moderate diagnostic sensitivity (30.0%, CI95 14.0–53.0%) for the molecular method. Mange-infected individually identified wolves showed a tendency for higher mortality versus uninfected wolves (ΔMortality 0.150, CI95 −0.165–0.458). Long-term serology data highlights the endemicity of sarcoptic mange in wild canids but uncovers multi-year epidemics. This study developed and evaluated a novel method for surveying sarcoptic mange in wildlife populations by the molecular detection of S. scabiei in faecal samples, which stands out for its high specificity and non-invasive character.

Newcastle disease virus transmission dynamics in wild peridomestic birds in the United Arab Emirates

To understand the dynamics of a pathogen in an animal population, one must assess how the infection status of individuals changes over time. With wild animals, this can be very challenging because individuals can be difficult to trap and sample, even more so since they are tested with imperfect diagnostic techniques. Multi-event capture-recapture models allow analysing longitudinal capture data of individuals whose infection status is assessed using imperfect tests. In this study, we used a two-year dataset from a longitudinal field study of peridomestic wild bird populations in the United Arab Emirates during which thousands of birds from various species were captured, sampled and tested for Newcastle disease virus exposure using a serological test. We developed a multi-event capture-recapture model to estimate important demographic and epidemiological parameters of the disease. The modelling outputs provided important insights into the understanding of Newcastle disease dynamics in peridomestics birds, which varies according to ecological and epidemiological parameters, and useful information in terms of surveillance strategies. To our knowledge, this study is the first attempt to model the dynamics of Newcastle disease in wild bird populations by combining longitudinal capture data and serological test results. Overall, it showcased that multi-event capture-recapture models represent a suitable method to analyse imperfect capture data and make reliable inferences on infectious disease dynamics in wild populations.

Multi-event capture-recapture analysis in Alpine chamois reveals contrasting responses to interspecific competition, within and between populations.

Understanding components of interspecific competition has long been a major goal in ecological studies. Classical models of competition typically consider equal responses of all individuals to the density of competitors, however responses may differ both among individuals from the same population, and between populations. Based on individual long-term monitoring of two chamois populations in sympatry with red deer, we built a multi-event capture-recapture model to assess how vital rates of the smaller chamois are affected by competition from the larger red deer. In both populations, mortality and breeding probabilities of female chamois depend on age and in most cases, breeding status the preceding year. Successful breeders always performed better the next year, indicating that some females are of high quality. In one population where there was high spatial overlap between the two species, the survival of old female chamois that were successful breeders the preceding year (high-quality) was negatively related to an index of red deer population size suggesting that they tend to skip reproduction instead of jeopardizing their own survival when the number of competitors increases. The breeding probability of young breeders (ages 2 and 3) was similarly affected by red deer population size. In contrast, in the second site with low spatial overlap between the two species, the vital rates of female chamois were not related to red deer population size. We provide evidence for population-specific responses to interspecific competition and more generally, for context-, age- and state-dependent effects of interspecific competition. Our results also suggest that the classical assumption of equal responses of all individuals to interspecific competition should be relaxed, and emphasize the need to move towards more mechanistic approaches to better understand how natural populations respond to changes in their environment.

Reproductive capacity of an endangered and recovering population of humpback whales in the Southern Hemisphere

Estimating demographic parameters is essential to assessing the recovery potential of severely depleted populations of marine mammal species such as the baleen whales, which were decimated by commercial whaling of the past century. The Oceania humpback whaleMegaptera novaeangliaepopulation is classified as endangered by the IUCN because of low numbers and a slow recovery rate. Nevertheless, an anomalously strong increase has recently been detected in the New Caledonia breeding population. To determine the drivers of population growth, reproductive parameters were estimated for the first time for a humpback whale population of Oceania. Based on an extensive monitoring program (1995-2018), recapture histories were reconstructed for 607 females and incorporated in multi-event capture-recapture models. As the females’ ages were generally unknown (87%), 2 models with contrasting age scenarios were investigated. For females of unknown age, the mature scenario assumed maturity at the first encounter, while the immature scenario assumed immaturity within 7 yr after the first encounter, unless the female was encountered breeding. These models respectively resulted in a calving interval of 1.49 yr (95% CI: 1.21-2.08) or 2.83 yr (95% CI: 2.28-3.56) and a calving rate of 0.67 or 0.35. The relatively high calving rate modelled by the mature model is consistent with high pregnancy rates recently observed in the migratory corridors of the Kermadec Islands and on the feeding grounds of the Antarctic Peninsula. Therefore, our results suggest that the recovery of the New Caledonia humpback whale population from past exploitation may be partially driven by an increased reproductive capacity.

Fitness of the pestiferous small rodent Mastomys natalensis in an agroecosystem in Mayuge district, Lake Victoria Crescent, Uganda

Abstract A 2.5-year study was conducted to understand the fitness of Mastomys natalensis in an agroecosystem in relationship with environmental predictors. The study was conducted in Mayuge district, in the Lake Victoria Crescent zone in Eastern Uganda. Fitness was measured in terms of survival, maturation and capture probability and estimated using multi-event capture-recapture models. Survival rates were higher after high rainfall in the previous month and increased with increasing population density of the animals. Maturation rate, on the other hand, showed no significant association with any predictor variables, while capture probability was significantly associated with sex of the animals, with higher capture probability for males. The results demonstrate that the fitness of M. natalensis in an agroecosystem is dependent on rainfall, sex and current population density. The aforementioned results were associated with increasing vegetation which provides cover for animal nesting and abundant food for the animals during rainfall periods and thus increased survival, high mobility in males in search for mates thus exposing animals to high chances of being captured and increased prey saturation at high population density resulting in high animal survival. These results have important implications for the timing of management strategies, i.e. control efforts should be enforced during the rainfall seasons to prevent high population buildup in the succeeding seasons.

Beak morphology predicts apparent survival of crossbills: due to selective survival or selective dispersal?

Dozens of morphologically differentiated populations, subspecies and species of crossbills (genus Loxia) exist. It has been suggested that this divergence is due to variation in the conifer cones that each population specialises upon, requiring a specific beak size to efficiently separate the cone scales. If so, apparent survival should depend on beak size. To test this hypothesis, we undertook multievent capture–recapture modelling for 6844 individuals monitored during 27 years in a Pyrenean common crossbill L. curvirostra population in a forest of mountain pine Pinus uncinata. Apparent survival was indeed related to beak width, resulting in stabilizing selection around an optimum that was close to the observed mean beak width, indicating that local crossbill beak morphology is adapted to the conifer they feed upon. Both natural selection (selective mortality) and selective emigration of maladapted individuals may explain our findings. As is often the case in capture–recapture analyses but rarely recognised, we could not formally decompose apparent survival into selective mortality versus selective permanent emigration. Nonetheless, there are several indications that selective permanent emigration should not be fully excluded. First, natural selection by itself would have to be unusually strong compared to other empirical estimates to create the observed pattern of apparent survival. Second, the observed mean beak width was a bit lower than the estimated optimum beak width. This can be explained by immigration of crossbills with smaller beaks originating from southern populations, which may subsequently have left the study area permanently in response to low food intake. This is in line with a detected transient effect in the data, yet apparently little influx from crossbills from northern Europe. When permanent emigration is phenotypically selective this will have ecological and evolutionary consequences, so this possibility deserves more attention in general.

Habitat‐driven life history variation in an amphibian metapopulation

Life‐history theory states that, during the lifetime of an individual, resources are allocated to either somatic maintenance or reproduction. Resource allocation tradeoffs determine the evolution and ecology of life‐history strategies and determine an organisms’ position along the fast–slow continuum. Theory predicts that environmental stochasticity is an important driver of resource allocation and therefore life‐history evolution. Highly stochastic environments are expected to increase uncertainty in reproductive success and select for iteroparity and a slowing down of the life history. To date, most empirical studies have used comparisons among species to examine these theoretical predictions. By contrast, few have investigated how environmental stochasticity affects life‐history strategies at the intraspecific level. In this study, we examined how variation in breeding site stochasticity (among‐year variability in pond volume and hydroperiod) promotes the co‐occurrence of different life‐history strategies in a spatially structured population, and determines life‐history position along the fast–slow continuum in the yellow‐bellied toad Bombina variegata. We collected mark–recapture data from a metapopulation and used multievent capture–recapture models to estimate survival, recruitment and breeding probabilities. We found higher survival and longer lifespans in populations inhabiting variable sites compared to those breeding in stable ones. In addition, probabilities of recruitment and skipping a breeding event were higher in variable sites. The temporal variance of survival and recruitment probabilities, as well as the probability to skip breeding, was higher in variable sites. Taken together, these findings indicate that populations breeding in variable sites experienced a slowing down of the life‐history. Our study thus revealed similarities in the macroevolutionary and microevolutionary processes shaping life‐history evolution.

Environmentally mediated reproductive success predicts breeding dispersal decisions in an early successional amphibian

Dispersal is a central mechanism in ecology and evolution. Dispersal evolution is driven by a trade-off between costs and benefits, which is influenced by interindividual variability and local environmental conditions (context-dependent dispersal). Many studies have investigated how dispersal decisions may be influenced by environmental factors, including density, predation and interspecific competition. Yet few have attempted to examine how habitat disturbance may affect the dispersal process in spatially structured populations. In early successional species, one might expect individuals to adjust their dispersal decisions based on two main factors that potentially have an influence on reproductive success: patch size and the level of patch disturbance. In this study, we examined how these two factors affect breeding success and dispersal decisions in an early successional amphibian, the yellow-bellied toad, Bombina variegata. To this end, we used capture–recapture data collected on a spatially structured population occupying 28 breeding patches. We took advantage of recent developments in multievent capture–recapture models to detect signs of context-dependent dispersal. The results revealed that the probability of successful reproduction and the number of newly metamorphosed individuals increased with both the size and the proportion of disturbance of a patch. In addition, our results showed that the factors affecting breeding success also influenced breeding dispersal probability. Large patch size negatively influenced emigration probability; in contrast, it positively influenced immigration probability. Equally, higher disturbance had a strong negative influence on emigration probability and slightly positively affected immigration probability. These findings strongly suggest that individuals make context-dependent dispersal decisions, adjusted to maximize future fitness prospects in a patch, allowing them to better cope with rapid changes in environmental conditions resulting from the ecological succession process. This opens new areas of potential research into the role of dispersal in organism specialization along an ecological succession gradient.

Use of hidden Markov capture-recapture models to estimate abundance in the presence of uncertainty: Application to the estimation of prevalence of hybrids in animal populations.

Estimating the relative abundance (prevalence) of different population segments is a key step in addressing fundamental research questions in ecology, evolution, and conservation. The raw percentage of individuals in the sample (naive prevalence) is generally used for this purpose, but it is likely to be subject to two main sources of bias. First, the detectability of individuals is ignored; second, classification errors may occur due to some inherent limits of the diagnostic methods. We developed a hidden Markov (also known as multievent) capture–recapture model to estimate prevalence in free‐ranging populations accounting for imperfect detectability and uncertainty in individual's classification. We carried out a simulation study to compare naive and model‐based estimates of prevalence and assess the performance of our model under different sampling scenarios. We then illustrate our method with a real‐world case study of estimating the prevalence of wolf (Canis lupus) and dog (Canis lupus familiaris) hybrids in a wolf population in northern Italy. We showed that the prevalence of hybrids could be estimated while accounting for both detectability and classification uncertainty. Model‐based prevalence consistently had better performance than naive prevalence in the presence of differential detectability and assignment probability and was unbiased for sampling scenarios with high detectability. We also showed that ignoring detectability and uncertainty in the wolf case study would lead to underestimating the prevalence of hybrids. Our results underline the importance of a model‐based approach to obtain unbiased estimates of prevalence of different population segments. Our model can be adapted to any taxa, and it can be used to estimate absolute abundance and prevalence in a variety of cases involving imperfect detection and uncertainty in classification of individuals (e.g., sex ratio, proportion of breeders, and prevalence of infected individuals).

A multi-event capture-recapture analysis of Toxoplasma gondii seroconversion dynamics in farm cats

BackgroundDomestic cats play a key role in the epidemiology of the parasite Toxoplasma gondii by excreting environmentally-resistant oocysts that may infect humans and other warm-blooded animals. The dynamics of Toxoplasma gondii seroconversion, used as a proxy for primo-infection dynamics, was investigated in five cat populations living on farms.MethodsSerological tests on blood samples from cats were performed every three months over a period of two years, for a total of 400 serological tests performed on 130 cats. Variations in seroconversion rates and associated factors were investigated using a multi-event capture-recapture modelling approach that explicitly accounted for uncertainties in cat age and serological status.ResultsSeroprevalence varied between farms, from 15 to 73%, suggesting differential exposure of cats to T. gondii. In farms with high exposure, cats could become infected before reaching the age of six months. Seroconversion rates varied from 0.42 to 0.96 seroconversions per cat per year and were higher in autumn and winter than in spring and summer.ConclusionOur results suggest inter-farm and seasonal variations in the risks of exposure to T. gondii oocysts for humans and livestock living on farms. The paper also discusses the role of young cats in the maintenance of environmental contamination by T. gondii oocysts on farms.

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season.

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long-lived seabird that experiences high bycatch rates in longline fisheries and strong population-level impacts due to this type of anthropogenic mortality. Analyzing a long-term dataset on individual monitoring, we compared adult survival (by means of multi-event capture-recapture models) among three close predator-free Mediterranean colonies of the species. Unexpectedly for a long-lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony-specific and mortality risk varies spatially.

Longitudinal survey of two serotine bat (Eptesicus serotinus) maternity colonies exposed to EBLV-1 (European Bat Lyssavirus type 1): Assessment of survival and serological status variations using capture-recapture models.

This study describes two longitudinal serological surveys of European Bat Lyssavirus type 1 (EBLV-1) antibodies in serotine bat (Eptesicus serotinus) maternity colonies located in the North-East of France. This species is currently considered as the main EBLV-1 reservoir. Multievent capture-recapture models were used to determine the factors influencing bat rabies transmission as this method accounts for imperfect detection and uncertainty in disease states. Considering the period of study, analyses revealed that survival and recapture probabilities were not affected by the serological status of individuals, confirming the capacity of bats to be exposed to lyssaviruses without dying. Five bats have been found with EBLV-1 RNA in the saliva at the start of the study, suggesting they were caught during virus excretion period. Among these bats, one was interestingly recaptured one year later and harbored a seropositive status. Along the survey, some others bats have been observed to both seroconvert (i.e. move from a negative to a positive serological status) and serorevert (i.e. move from a positive to a negative serological status). Peak of seroprevalence reached 34% and 70% in site A and B respectively. On one of the 2 sites, global decrease of seroprevalence was observed all along the study period nuanced by oscillation intervals of approximately 2–3 years supporting the oscillation infection dynamics hypothesized during a previous EBLV-1 study in a Myotis myotis colony. Seroprevalence were affected by significantly higher seroprevalence in summer than in spring. The maximum time observed between successive positive serological statuses of a bat demonstrated the potential persistence of neutralizing antibodies for at least 4 years. At last, EBLV-1 serological status transitions have been shown driven by age category with higher seroreversion frequencies in adults than in juvenile. Juveniles and female adults seemed indeed acting as distinct drivers of the rabies virus dynamics, hypothesis have been addressed but their exact role in the EBLV-1 transmission still need to be specified.

Insights on dispersal and recruitment paradigms: sex- and age-dependent variations in a nomadic breeder.

Sex- and age-dependence in recruitment and dispersal are often explained by costs arising from competition for holding a breeding territory over the years-a typical feature of species living in stable habitats. For instance, long-lived birds with male territoriality often exhibit large variation in recruitment age and higher dispersal in females and young individuals. As a corollary, we expected that species with ephemeral habitat suitability, and hence nomadic breeding, would show weak age- and sex-dependence in dispersal and low variation in recruitment age, because territory ownership is not maintained over the years. In addition, the higher cost of reproduction in females might not be (over)compensated for by costs of territoriality in males. Accordingly, females would recruit later than males. We explored these variations using multievent capture-recapture models over 13years, 3479 (2392 sexed) slender-billed gulls (Chroicocephalus genei) and 45 colony sites along the French Mediterranean coast. As expected, variability in recruitment age was low with males recruiting earlier than females. Nonetheless, dispersal in and out of the study area decreased with age and was slightly higher in males than in females. Decreased dispersal with age might result from foraging benefits associated with increased spatial familiarity. Higher dispersal in males might be explained by a male-biased sex ratio or higher philopatry benefits in females (arising from their higher cost of reproduction). Sex- and age-dependent dispersal and recruitment may thus occur in the absence of year-to-year breeding territory ownership, which stresses the importance of considering other processes in shaping recruitment and dispersal patterns.

Disentangling the effects of predation and oceanographic fluctuations in the mortality of two allopatric seabird populations

AbstractLife‐history traits of migratory seabirds are influenced by changing conditions at breeding and wintering grounds. Climatic conditions and predation are known to impact populations’ survival rates, but few studies examine their effect simultaneously. We used multievent capture–recapture models to assess mortality due to environmental conditions and predation in breeding European storm petrels (Hydrobates pelagicus) in two allopatric colonies (Mediterranean and Atlantic). Predatory mortality at the colonies showed annual variation, being around 0.05 in certain years. Mortality at sea differed between the two oceanic basins, and was lower in the Mediterranean colony [0.11, 95% CI (0.09, 0.14)] when compared to the Atlantic colony [0.18, 95% CI (0.15, 0.22)]. The Western Mediterranean Oscillation index (WeMOi) explained 57% of the temporal variability in mortality of Mediterranean breeders. In comparison, 43% of the temporal variability in mortality of Atlantic breeders was explained by the winter St Helena index (wHIX) and El Niño‐Southern Oscillation index (wENSO). Our results suggest that Mediterranean breeders remain in this basin for wintering where they may face lower migratory costs and more favourable environmental conditions. In contrast, Atlantic breeders’ mortality may be due to higher cost of migration, changing upwelling conditions in the Benguela current and heavy storms over their migratory route during La Niña events. This study underlines the importance of modelling separately different causes of mortality when testing the effects of climatic covariates.

Varying demographic impacts of different fisheries on three Mediterranean seabird species.

Fisheries have an enormous economic importance, but reconciling their socio-economic features with the conservation and sustainability of marine ecosystems presents major challenges. Bycatch mortality from fisheries is clearly among the most serious global threats for marine ecosystems, affecting a wide range of top predators. Recent estimates report ca. 200,000 seabirds killed annually by bycatch in European waters. However, there is an urgent need to rigorously estimate actual mortality rates and quantify effects of bycatch on populations. The Mediterranean Sea is one of the most impacted regions. Here, we estimate for the first time both bycatch mortality rates and their population-level effects on three endemic and vulnerable Mediterranean taxa: Scopoli's shearwater, Mediterranean shag, and Audouin's gull, that die in different types of fishing gears: longlines, gillnets andsport trolling, respectively. We use multi-event capture-recapture modelling to estimate crucial demographic parameters, including the probabilities of dying in different fishing gears. We then build stochastic demography models to forecast the viability of the populations under different management scenarios. Longline bycatch was particularly severe for adults of Scopoli's shearwaters and Audouin's gulls (ca. 28% and 23% of total mortality, respectively) and also for immature gulls (ca. 90% of mortality). Gillnets had a lower impact, but were still responsible for ca. 9% of juvenile mortality on shags, whereas sport trolling only slightly influenced total mortality in gulls. Bycatch mortality has high population-level impacts in all three species, with shearwaters having the highest extinction risk under current mortality rates. Different life-history traits and compensatory demographic mechanisms between the three species are probably influencing the different bycatch impact: for shearwaters, urgent conservation actions are required to ensure the viability of their populations. Results will be very useful for guiding future seabird conservation policies and moving towards an ecosystem-based approach to sustainable fisheries management.

Demography of the critically endangered Balearic shearwater: the impact of fisheries and time to extinction

Summary World‐wide, many seabirds are affected by fisheries in opposing ways: as a source of mortality from bycatch, but also by providing discards as a predictable and abundant food resource. This applies to the Balearic shearwater Puffinus mauretanicus, the most endangered European seabird, whose time to extinction was estimated at only ˜40 years a decade ago. Since the previous assessment, new data and more sophisticated demographic modelling have become available, and new fishing policies from the European Union (Common Fisheries Policy, CFP) will apply, posing different scenarios for the viability of the species. Thus, there is both an urgent need and an opportunity for a more reliable update of the conservation status of the species. Demographic data were collected between 1985 and 2014 at one of the world's largest colonies. Most demographic parameters were estimated using multievent capture–recapture modelling. Some parameters, such as bycatch rate, immature individual survival and recruitment, were estimated for the first time. We incorporated estimates into stochastic population models to forecast time to extinction and assess the viability under different management scenarios, accounting for upcoming fishing policies. Adult survival was much lower than expected (0·809, SE: 0·013) and largely influenced by bycatch, which accounted for a minimum of 0·455 (SE: 0·230) of total mortality. Breeding success was positively correlated with discard availability. Recruitment started at low rates in 3‐year‐old birds (0·030, SE: 0·0455), increasing in following age classes and was almost complete at 6 years. Under the present scenario, we predict a time to extinction of 61 years (95% CI: 55–69). Synthesis and applications. Population projections suggest that the actual impact of fisheries on Balearic shearwaters is unsustainable and the imminent discard ban under the new Common Fisheries Policy may accelerate the declining trend. This study demonstrates that reducing the bycatch rates of fisheries is an unavoidable and urgent conservation measure for avoiding the extinction of the species. We also advise setting up demographic long‐term studies, to allow researchers to diagnose, with reliability, the effectiveness of management actions. These actions will also benefit many other marine top‐predator species affected by this anthropogenic impact.

Modelling survival and breeding dispersal to unobservable nest sites

Context Demographic parameters in wildlife populations are typically estimated by monitoring a limited number of individuals in observable sites and assuming that these are representative of the whole population. If individuals permanently disperse to unobservable breeding sites, recruitment and immature survival are expected to be negatively biased and breeding-site fidelity cannot be investigated. Aims To develop a method to obtain unbiased estimated of survival, recruitment and breeding dispersal when individuals can move to, or recruit in, unobservable sites. Methods We used the flexibility of multi-event capture–recapture models to estimate dispersal and recruitment to unobservable sites, merging observations made at two sites within the same breeding locations. We illustrated the model with data on little penguin (Eudyptula minor) breeding in artificial as well as in natural nests. Natural nests are unknown or inaccessible and birds in these sites remain unobservable. Encounters at beaches surrounding the colony suggested that marked animals can permanently move to unobservable nests. We built the multi-event model considering two possible states of the individuals (alive breeding in a nest box and alive in a natural nest) and three types of observations (encountered at a nest only, encountered at the beach only and encountered at both places). This model ensured that the breeding dispersal to unobservable places became estimable. Key results Results indicate that the estimated survival was 8% higher than when recaptures at artificial nests were analysed alone. Also, fidelity to artificial nests was 12% lower than to natural nests. This might reflect the greater availability of natural sites or, alternatively, a heterogeneity between these two types of nest. Conclusions We obtained an estimate of local survival of little penguins breeding at Penguin Island that incorporates the permanent migration to unobservable sites and found an asymmetric dispersion towards natural nests. Implication Our conclusions suggest a need for more careful treatment of data derived from artificial sites alone, as demographic parameters might be underestimated if animals prefer natural breeding sites or if they are in greater proportion compared with artificial ones. The analytical approach presented can be applied to many biological systems, when animals might move into inaccessible or unobservable breeding sites.