Population Ecology

Demographic models are crucial for uncovering the mechanisms underlying changing populations trajectories and structure and to identify the drivers of such changes. Common pochard populations of North east/North west Europe have experienced a sharp decline accompanied by an increase in male proportions among adults since the mid-1990. We used a two-sex, two-stage deterministic matrix population model, to perform a prospective perturbation analysis and to explore, through simulations, some plausible causes underlying the observed decline and change in sex structure. We show that Common pochard populations are more sensitive to changing survival than to changes in productivitys components (clutch size, nest survival...). However, due to an environmental variance much higher than that of survival, components of productivity, especially nest survival, would be the main drivers of Common pochard populations growth rate, a finding supported by empirical data. More importantly, we show that although sex-specific changes in survival at any stage of the life cycle are potent drivers of both population growth rate and changing sex ratio, there is no need to resort to them for explaining the increasing proportions of males such as observed in Common pochard. Because adult males display higher survival than adult females (0.74 against 0.64 on average), any factor affecting recruitment (nest or first year survival) increases the weight of adults into the populations and hence the proportion of males. Thus, in species displaying sex-biased mortality, such as many ducks, decreasing recruitment can underly declining population size and changes in sex structure at the same time, emphasising the importance of accounting for males in monitoring schemes and demographic models.

Arctic and Boreal raptor communities will continue to be affected by borealization and other climate change related processes, providing a challenge for ecologists predicting future sates. However, by using community assembly theory and species traits, future communities may be predictable. In this study, we analyzed variation in reproduction traits as a consequence of diet specialization for 29 raptors, 2 skuas and 3 corvids. We assessed and implemented foraging traits for specialists and generalists into predator-prey models from which successful invasion conditions were derived. Specialist raptors produced larger clutch sizes, had a higher proportion of fledged per clutch and also expressed more variation compared to generalist raptors. These results suggest a relationship between diet specialization and reproductive traits which was also observed within phylogenetic orders. Specialist owls (Strigiformes) produced higher clutch sizes with a larger clutch range compared to generalist owls. The same pattern was observed for falcons (Falconiformes). No clear difference in reproduction was observed for specialist and generalist hawks, kites and eagles (Accipitriformes). Corvids expressed clutch sizes similar to that of specialist raptors while having the lowest proportion of fledged per clutch. Differences in foraging traits between specialists and generalists could be distinguished using functional response curves. A predator-prey model parameterized with foraging trait data showed that a generalist can coexist with a resident specialist if it has access to prey unavailable to the resident specialist. Otherwise, the native specialist outcompetes the invading generalist due to foraging efficiency. The combined empirical and theoretical findings in this study show how diet specialization affects both reproduction and the potential invasion success of raptors.

Age-dependent survival is central to understanding population dynamics and life-history evolution. We analysed carcass weight and age-at-harvest data from 6022 red foxes (Vulpes vulpes) collected across Sweden between 1967 and 1971 to evaluate latitudinal effects on body mass and age-dependent survival. Carcass weights decreased from south to north in both adults and sub-adults, contrary to Bergmann's rule, with southern foxes weighing approximately 1.27 times more than northern foxes. The latitudinal weight gradient exceeded the sex difference in both age classes, and no sex x region interaction was detected. The decrease in weight with latitude is consistent with reduced prey availability and harsher winter conditions in the north, which limit growth and body size during development. Using a Bayesian age-at-harvest model with region-specific population growth rates (lambda), we estimated age-dependent survival probabilities for four latitudinal regions and both sexes. Despite the strong latitudinal gradient in weight, survival did not show a corresponding pattern - regional differences were uncertain, with all credible intervals spanning zero. Regional population growth rates were consistent with slight decline in the north and near-stability in the south-central region, which suggests that body condition and population dynamics are coupled at the regional scale despite no survival gradient. The decoupling of body condition and survival across regions suggests that mortality patterns are similar across the latitudinal gradient. We discuss these patterns in terms of latitudinal productivity gradients, prey availability, and life-history trade-offs in a widely distributed carnivore.

Managers must make decisions in the face of uncertainty, especially when available resources are limiting. Identifying thresholds when certain conditions are met or exceeded enable the potential to mitigate risks. In 2005, sustainable harvest levels of willow ptarmigan were identified to avoid harvest efforts exceeding three hunter days km2. Here we evaluate these recommendations by analyzing line transect counts and harvest data from six areas forming three open/closed pairs in a region of state managed willow ptarmigan harvest. We developed three sets of Bayesian hierarchical models, one static distance model, and two dynamics models. One mechanistic hazard model and a Gompertz phenomenological model. Adult and juvenile density showed pronounced year-to-year variation that was largely synchronous across all six sites regardless of hunting status. The harvest effort parameter shows a striking difference between the two models. In the Hazard model, is positive, and excludes zero with near certainty, but in the Gompertz model, the parameter is highly uncertain. However, the two models do not contradict each other but answer complementary questions with different sensitivity to the harvest signal, harvest mortality is additive at the individual level, but this additive mortality is masked at the level of population abundance. The demographic cost of harvest is therefore real and quantifiable through the survival chain, but bounded in the long run by the stabilizing dynamics. A fixed limit anchored to monitored effort and bag is not a crude substitute for adaptive management but the appropriate design under the information commonly at hand. It will be a precautionary instrument grounded in the one relationship this study establishes firmly, the translation of hunter effort into harvest mortality.

O_LIUnderstanding the processes underlying successful eradication of invasive species is essential for achieving global island conservation goals. Despite the widespread availability of capture records from eradication programs, modeling frameworks that utilize these datasets to elucidate spatio-temporal population dynamics remain underdeveloped. C_LIO_LIIn this study, we reconstructed the spatio-temporal population dynamics of the small Indian mongoose on Amami-Oshima Island (712 km{superscript 2}), Japan, where the species was introduced in 1979 and officially declared eradicated in 2024 after more than 30 years of systematic removal. We integrated introduction records, capture data, and monitoring data using a hierarchical harvest-based model (HBM). To evaluate the models capacity to support management decisions and assess eradication success, we conducted retrospective analyses and compared estimated eradication probabilities with those obtained from a rapid eradication assessment (REA; Samaniego-Herrera et al., 2013). C_LIO_LIThe estimated population size (before reproduction) peaked at 5,449 individuals (95% CI: 4,703, 6,175) in 2000 and subsequently declined almost monotonically. The maximum invaded area was 547.78 km{superscript 2} (posterior median, 95% CI: 496.47, 566.04) in 2009, indicating that the removal program successfully prevented island-wide expansion. Retrospective analyses showed that population estimates remained within the 95% credible intervals of the full dataset estimates, demonstrating temporal consistency. Eradication probabilities estimated by the HBM were substantially higher than those from the REA, highlighting the sensitivity of estimates to fine-scale heterogeneity in detection processes. C_LIO_LISynthesis and applications: Hierarchical HBMs provide a powerful framework for reconstructing, predicting, and evaluating invasive species eradication dynamics. Being aware of the limitations for application to eradication evaluations, HBMs can support adaptive management in long-term eradication programs and improve our understanding of the mechanisms underlying successful eradication. C_LI

Mortality causes and survival rate estimates from individual follow-ups of GPS-collared animals provide reliable demographic data, but such estimates are rare for red foxes Vulpes vulpes. We used data from 126 GPS-collared red foxes tracked between 2011 and 2019 across a latitudinal gradient in central Sweden and Norway to quantify mortality causes and estimate annual survival probabilities using the Andersen-Gill extension of the Cox proportional hazards model. Hunting was the dominant mortality cause (63%), followed by vehicle collisions (14%), stress or malnutrition (11%), sarcoptic mange (9%), and predation (3%). Annual survival was 0.58 for adults and 0.32 for subadults. Subadults had approximately twice the hazard of adults, males had elevated hazard relative to females, and mortality risk was highest in autumn and early winter. These collar-based estimates are strikingly similar to mark-recovery estimates from the same region nearly five decades earlier, despite an intervening population collapse from sarcoptic mange, subsequent full recovery, and a major decline in harvest pressure. This convergence is consistent with density-dependent regulation at a food-determined and food-limited carrying capacity, suggesting that current harvest levels are not limiting the population. Furthermore, human-caused mortality was approximately four times greater than natural mortality, illustrating that living in close proximity to humans -- which often favours generalist predators such as red foxes -- may also come at a cost of increased mortality risk. Developing validated census methods for red foxes in boreal Scandinavia is identified as a key priority for quantitative population management.

As anthropogenic threats have intensified over the past 500 years, we find ourselves in the midst of a sixth mass extinction, with continued losses of biodiversity threatening ecosystem stability. This biodiversity loss has caused species extinctions across taxa, and placed several others at high risk of functional extinction. These disturbance-driven impacts represent one of the most acute biodiversity crises facing global marine systems. Species exhibiting slow life histories characteristically have low resilience to disturbance. Here, we assess the risk of functional extinction and identify policy pathways for population recovery of the slow-living, Critically Endangered elasmobranch, the spinetail devil ray (Mobula mobular). We develop a stochastic, state-structured Integral Projection Model (IPM) parameterised with demographic data collected from fishery landings data in India, the world's largest mobulid fishery, and supplemented with data on vital rates from published literature. Using the IPM, we estimate that the population is declining at approximately 12% annually, experiencing substantial limiting pressure from fisheries overexploitation and failing to approach its biological maximum growth potential. Our results indicate that populations of M. mobular will be at high risk of functional extinction if 'business as usual' harvest scenario persists for another decade. We further show that long-term population recovery is only possible if survival increases significantly across all size classes, especially among large reproductive females, alongside a concurrent increase in fecundity. We conclude that no single policy measure is sufficient to recover population of M. mobular along the southeastern coast of India. Instead, combined protection through maximum bycatch mitigation and protection of nursery areas in no-take zones will be required for population recovery. This research demonstrates that recovery of overexploited populations often requires integrated resource management across life stages, and that the Critically Endangered M. mobular warrants urgent conservation action to avoid functional extinction.

Identifying optimal grazing intensities for sustainable population growth is crucial for informing management strategies. Community-level studies frequently find that biodiversity peaks at intermediate grazing intensities, known as the Intermediate Disturbance Hypothesis. However, whether this hypothesis applies to population-level performance remains untested. Our stochastic integral projection models, parameterized with five-year demographic data of two co-occurring species, Morina chinensis and Deyeuxia flavens, on the Tibetan Plateau grasslands., indeed show a hump-shaped response in stochastic population growth rate ({lambda}S) to grazing intensities, providing empiral support for the Intermediate Disturbance Hypothesis at population level. Furthermore, populations with demographic compensation among vital rates are better able to buffer temporal variation in annual population growth rate and exhibit a much smaller decline in {lambda}S under heavy grazing. Our study provides mechanistic insights into demographic processes driving population dynamics across grazing levels, thereby better informing grazing management strategies.

Diaphorina citri is the primary vector of Huanglongbing (HLB), a devastating disease that affects global citrus production. Effective biological control using the parasitoid Tamarixia radiata represents a sustainable alternative to chemical insecticides, but its efficacy depends heavily on environmental variables and resource availability. In this work, we develop a four-population mathematical model to analyze the dynamics of D. citri in the presence of T. radiata, a known parasitoid. Our model incorporates the cyclical and periodic behavior of citrus flushing (new shoots), providing a realistic representation of resource-limited dynamics as observed in field conditions. Through comparative simulations of three scenarios, without parasitoid, a single initial introduction, and periodic augmentative releases, we characterize and compare the impact of T. radiata as a biological control agent. Our results show that periodic releases maintain pest suppression, whereas a single introduction only delays pest recovery. By aligning theoretical modeling with ecological reality, this framework supports the role of T. radiata in pest suppression and provides a baseline for future work on optimal and cost-effective release strategies.

Environmental variability can strongly alter coexistence among competing species and their extinction risk, particularly when population dynamics are shaped by behavioral interactions, such as fear. In this work, we develop a novel stochastic differential equation competition model that incorporates both non-consumptive fear effects and environmental variability to investigate how behavioral interactions influence species coexistence under random fluctuations. Our result reveals that environmental stochasticity can drive species to extinction even when the corresponding deterministic system admits coexistence. In particular, under an explicit stability condition on the fear and competition parameters and sufficiently strong averaged noise intensities, we prove that both competing species become extinct exponentially almost surely. Conversely, we derive a stochastic persistence criterion in terms of fear, competition, and noise-induced suppression parameters for the fearful species. We further demonstrate that environmental noise may reverse classical competition-exclusion outcomes, leading to qualitatively different long-term dynamics from those predicted deterministically. These results provide rigorous thresholds separating stochastic extinction from persistence and highlight the critical role of environmental variability in fear-mediated competitive ecosystems. From an applied perspective, these results provide insight into how behavioral interactions and environmental variability influence species survival, with potential applications in ecological management and conservation.

Many Sockeye salmon (Oncorhynchus nerka) populations have declined over recent decades, and climate change is likely to exacerbate these declines through direct and indirect ecological effects. The response to the associated environmental changes is likely to vary among life stages, populations, and regions. Quantitative estimates of climate change driven impacts that account for this variability could fill a critical gap and provide forward-looking insights into how sockeye are expected to respond to future climate-driven change across their lifecycle. To address this need we developed a hierarchical population dynamics model parameterized with juvenile, adult return and spawner abundance data from 13 sockeye salmon populations from Washington State to northern British Columbia. We used a formal causal inference framework that paired salmon abundance data with a suite of environmental covariates hypothesized to represent ecological conditions across the lifecycle. We used the model to estimate population-specific responses to each environmental driver, then combined parameter estimates with projections from down-scaled climate change models to estimate productivity responses to anticipated environmental change. We found that historical sockeye productivity was strongly associated with environmental covariates, which explained more interannual variability in return abundance than spawner abundance in most populations. However, the life stages and specific environmental covariates with the largest impacts differed among populations and regions, often displaying a latitudinal gradient. Increases in coastal ocean temperatures and mixed layer depth generally had negative effects though they varied among regions. Increased freshwater summer rearing and return migration temperatures had weaker but consistently negative effects. Under future climate conditions, projected changes in these environmental covariates are expected to result in substantial declines in productivity across most populations. Sockeye salmon display varying degrees of sensitivity to climate change across life stages, populations, and regions. Effective future management will require explicitly accounting for these life stage and population-specific responses.

Inbreeding depression (the reduction in fitness associated with inbreeding) has been demonstrated in a wide range of animals, but despite its ubiquity, is not an inevitable consequence of inbreeding. As a result, there is uncertainty about the extent to which inbreeding depression poses an ongoing risk to endangered species currently experiencing significant demographic recovery. Quantifying inbreeding depression will be critical if we want to understand these risks. A comprehensive quantification of the fitness costs of inbreeding requires detailed individual-based longitudinal data so lifetime impacts can be assessed. Here, we use an extraordinarily detailed long-term dataset on Mauritius kestrels (Falco punctatus) to explore inbreeding depression in a population currently experiencing significant demographic recovery. To do so, we constructed a social pedigree of 1,758 individuals and combined this with 1,240 nest records and 1,411 individual resighting histories to explore lifetime fitness effects over a 30-year period. Inbreeding increased significantly over time as the population recovered before stabilising. Inbred eggs were less likely to survive to fledging. Inbred adult male and female birds had significantly lower annual reproductive success than outbred individuals because of a lower annual egg-to-fledgling survival probability. This resulted in significantly lower lifetime reproductive success in inbred females but not males, which showed a negative trend. Population growth was negative and extinction risk increased slightly at current levels of inbreeding. These impacts will become more severe should inbreeding levels increase in the future, which is highly likely given ongoing genomic erosion. Taken together, our results demonstrate significant fitness costs associated with inbreeding in Mauritius kestrels, which pose an ongoing risk to population viability. This suggests that monitoring and managing inbreeding risks in endangered species will likely be required even in populations that are showing significant demographic recovery in response to conservation interventions.

Agroecosystems are dynamic ecosystems, constituted by patches of vegetation and agricultural use, where biodiversity is shaped by spatial and temporal variability. While most studies have focused on spatial composition and configuration, the role of temporal variability remains poorly understood. Yet, temporal dynamics can strongly modify species composition, abundance, and persistence in ecological communities. Temporal variability is particularly relevant in agroecosystems with rainfed agriculture where environmental conditions shift dramatically between rainy and dry seasons. In this paper, we assess the role of temporal variability on biodiversity maintenance in an agricultural matrix using a metacommunity model that simulates an agricultural landscape under rainfed conditions, that is, with abrupt seasonal changes in the agricultural patches. This model couples a local community network dynamic with a migration dynamic and is based on empirically documented features of rainfed agricultural matrices. Our results show that temporal variability provides new opportunities for species to recover from low densities. However, the effect of temporal variability is not straightforward. It depends on the initial and final conditions, the migration and mortality rates and the intensity of temporal variability. Overall, our findings highlight the need to further investigate temporal variability to better understand its role in shaping biodiversity in agricultural landscapes.

Space-for-time Cormack-Jolly-Seber (CJS) models have been developed to estimate survival of migrating animals that are imperfectly detected at spatially discrete sampling stations. However, space-for-time CJS models typically ignore survival over time and age-specific probability of movement, missing the diversity of life history strategies. To address these limitations, we extended space-for-time Cormack-Jolly-Seber models to explicitly incorporate time, age, and individual-level covariates to account for diverse life history strategies. We incorporate a sub-model that includes uncertainty in individuals age using a finite-mixture model. The detection and transition probabilities were parameterized using generalized linear models, facilitating flexible model specification and inclusion of spatiotemporal and individual covariates. We apply the model to detections of juvenile steelhead (Oncorhynchus mykiss) from two populations in the Snake River Basin in Idaho, USA, to estimate trends in survival with respect to population, age, time, and the length of individuals. Additionally, we generalized the model so that it can be applied to other systems and populations with different life history strategies and monitoring infrastructure.

The moonfish Mene maculata (Bloch and Schneider, 1801), the sole extant species of the family Menidae, is widely distributed in the Indo-West Pacific. Although its spawning season has previously been inferred from seasonal variation in the gonadosomatic index (GSI), the histological basis for interpreting GSI as an indicator of spawning-capable status remains limited. Here we describe the gonadal structure of male and female M. maculata and evaluate how GSI and standard length relate to spawning-capable status based on germ cell development. The testis was lobular and exhibited an unrestricted spermatogonial distribution, a structure widely observed among neoteleosts, indicating that the phylogenetic distinctiveness of M. maculata was not associated with a distinctive testicular structure. In the spawning-capable female, oocytes at multiple developmental stages, from primary growth to oocyte maturation, co-occurred within the ovary, indicating asynchronous ovarian development. This finding suggests that M. maculata may be a batch spawner rather than a total spawner as previously inferred. The spawning-capable female had a GSI consistent with previously inferred spawning season estimates. By contrast, histologically examined males were spawning capable at GSI values lower than those previously associated with the inferred spawning season, suggesting that male spawning-capable status may persist beyond the period inferred from elevated GSI alone. This study provides the first histological description of reproductive biology in the phylogenetically distinctive M. maculata and establishes a histological basis for interpreting GSI as a reproductive indicator.

Humans are efficient and deadly predators, yet they may also interact with wildlife in non-lethal ways. This study explores how interactions with lethal and non-lethal human "superpredators" alter predator-prey dynamics using an agent-based modelling approach. Our model incorporates both the consumptive (lethal) and non-consumptive (behavioural) effects of humans, as well as of predators on prey. We explored how the replacement of apex predators by humans affects mesopredator-prey dynamics, with particular emphasis on trophic targeting and differences between lethal and non-lethal interactions. We found that human superpredators have a greater effect on model outcomes than apex predators. When superpredators consume mesopredators alone or with prey, the probability of mesopredator-prey coexistence increases to a greater extent than when apex predators consume mesopredators. In contrast, superpredators consuming only prey slightly increases overall extinction risks and reduces coexistence. Non-lethal superpredators, despite eliciting anti-predator responses in mesopredators and prey, had a negligible effect on population dynamics. Our findings demonstrate that human superpredators may functionally replace apex predators when they are lethal. However, non-lethal interactions with humans may not be as ecologically significant as lethal interactions, even when humans induce anti-predator responses.

Climate change is expected to alter plant populations not only through direct environmental shifts but also via changes in biotic interactions, such as with herbivores and pathogens. As plant species are also expected to differ in their responses to both climate and antagonists, plant responses to both factors are expected to be variable and species-specific. To assess whether interactive effects of climate and antagonists on plant population dynamics are common and whether the strength and direction of plant responses vary across species, we conducted a multi-year field experiment that manipulated realistic climate change and experimentally reduced insect herbivores and fungal pathogens. We measured responses of plant vital rates, such as survivorship, growth, and reproduction across six grassland species. Using Integral Projection Models (IPMs) and Life Table Response Experiments (LTREs), we quantified changes in population growth rate across experimental treatments and the contribution of each vital rate to that observed change. Two of the study species declined so drastically over the course of the experiment that demographic quantification of population growth rates was not possible. From the remaining species, Bromus erectus and Plantago lanceolata show significant interactive responses of climate and antagonist reduction on population growth rates. In contrast, Dianthus carthusianorum and Tragopogon orientalis showed limited responses to experimental treatments. Notably, our results indicate that in some species biotic interactions may amplify the effects of climate change: the presence of plant antagonists exacerbates the negative effects of the future climate treatment on plant population dynamics. Our findings highlight the complexity in predicting plant population responses to climate change and provide insights for grassland management under future environmental conditions.

ContextThe movement of vertebrate hosts across urbanized landscapes can play a key role in the transmission of direct-contact diseases. Understanding how wildlife hosts move in urban landscapes, and how transmission is affected by their landscape-constrained and disease-altered movements, is imperative for better predicting the spread of disease. ObjectiveWe assess how the movement of red foxes (Vulpes vulpes) according to landcover type, and their infection status, affect the spread of mange (caused by Sarcoptes scabiei) in an urbanized landscape. MethodsWe developed a mange transmission model (MTM) using an agent-based model to compare two movement behaviours of foxes in Scarborough (Ontario, Canada): random and landcover-based. We further assessed the effects of movement on disease transmission by considering the foxs infection status and comparing a range of movement probability scenarios. We quantified the number of effective contact events and the effective reproduction number (Re) according to each scenario. ResultsWe found that both landcover-dependent movement and infection status influenced the spread of mange within fox populations. The number of effective contact events and effective reproduction number Re was greatest when landscape heterogeneity was included in the model and foxes moved through paths of least resistance to movement, and when susceptible and infected foxes had an equal probability of leaving a fragmented habitat patch. ConclusionsOur findings suggest that mange spread may be accelerated along movement corridors in fragmented, heterogenous landscapes. As urban areas expand and remnant habitat within these is further lost and animals are relegated to fewer movement pathways, disease transmission may increase.

Movement models often assume that animal decisions depend on perceived environmental conditions. However, optimal foraging and cognition theory predict that accumulated experiences should drive changes in animal motivation and decision-making. Here, we propose a mechanistic framework that models state-switching habitat selection as a history-dependent process, where internal states, such as fear or energy, emerge as latent variables accumulating through environmental exposure. Simulations showed that our model accurately recovers memory timescales and cumulative effects of environmental exposure on behavioural switching that would be undetected by existing state-switching habitat selection models. Applied to woodland caribou, it reveals that an individual integrates predation risk experienced over the previous 15 days, but food intake over only 2 days, when deciding to remain or leave an area. Our model advances perspectives on movement ecology by quantifying how accumulated experiences shape changes in motivation driving animal movement decisions.

Long-lived plants facing anthropogenic disturbance often exhibit recruitment failure despite persistent reproductive adults, generating extinction debt masked by longevity. However, whether adult presence reliably reflects environmental suitability for recruitment remains unclear. Here, we examine ontogenetic niche differentiation and its consequences for recruitment in Jubaea chilensis, an endangered long-lived Mediterranean palm with an aging population. We assigned individuals within the largest known population to four ontogenetic stages and characterized their environmental niches using climatic, edaphic, topographic, and vegetation variables. We then applied spatial and multivariate analyses, including Random Forest models to evaluate environmental segregation and identify predictors of seedling establishment. Age classes occupied significantly different environmental niches, with the greatest differentiation between seedlings and reproductive adults. Saplings and adult differentiation reflected mainly topographic variables at landscape scale, whereas seedling establishment was primarily predicted by microhabitat conditions (vegetation cover heterogeneity, east-facing slope orientation, and soil texture). This pattern is consistent with niche reconfiguring throughout the life cycle, suggesting that adult occurrence and recruitment suitability respond to distinct environmental conditions. Over one-fifth of sampled individuals occupied high-suitability sites without recruitment, suggesting that ontogenetic niche shifts are associated with a spatial decoupling between adult persistence and recruitment, consistent with demographic collapse independent of habitat degradation. This failure is likely mediated by insufficient effective seed dispersal, as the sole disperser (Octodon degus) preys on most seeds before dispersal. Conservation strategies based solely on adult distribution may therefore overestimate effective habitat and underestimate extinction risk in long-lived species.