Current wolf recolonisation is fuelling polarised debates over nature restoration versus the preservation of rural traditions. Understanding how public attitudes evolve as wolves return and how acceptance coincides with suitable wolf habitat is key to fostering coexistence. We analysed nationwide attitude surveys from early and advanced stages of recolonisation in Switzerland (2015 and 2023), prior to the recent shift towards proactive lethal management. First, we modelled the spatial distribution of attitudes towards wolves, measured on a 5-point Likert scale from support to opposition, to identify spatiotemporal changes across the country. Second, we examined how attitudes and their changes related to respondents’ surroundings, including exposure to wolves and livestock depredation, human demographic, and socioeconomic factors. Third, we compared acceptance with ecological habitat suitability to locate areas of potential socioecological conflict. Despite wolf range expansion, nationwide acceptance of wolves in Switzerland remained stable between 2015 and 2023. We consistently found more negative attitudes in mountain regions than in lowlands, driven more by socio-demographic factors than by exposure to wolves. However, local attitude shifts were highly variable and mostly linked to the respondents’ geographical location and their spatial proximity to each other. This suggests that changes in attitudes were influenced more by community dynamics than by ecological factors related to recolonisation or conflicts. Our findings highlight the importance of a nuanced approach to wildlife management that recognises areas of potential socioecological conflict as priorities for conservation, and of promoting societal discourse on community beliefs and values in landscapes of contested wolf presence.

Current wolf recolonisation is fuelling polarised debates over nature restoration versus the preservation of rural traditions. Understanding how public acceptance evolves as wolves return and how it coincides with suitable wolf habitat is key to fostering coexistence. We analysed nationwide surveys from early (2015) and advanced (2023) stages of wolf recolonisation in Switzerland. First, we modelled the spatial distribution of public acceptance of wolves, measured on a 5-point Likert scale from support to opposition, to determine how acceptance related to respondents' local environments, including exposure to wolves and livestock depredation, as well as geographical, human demographic, and socioeconomic factors. Second, we mapped changes in acceptance between the two survey years and examined how they related to changes in exposure to wolves and livestock depredation. Third, we spatially compared acceptance with ecological habitat suitability to locate areas of potential socioecological conflict. Despite wolf range expansion and population growth, nationwide acceptance of wolves in Switzerland remained stable between 2015 and 2023. We found consistently lower acceptance in mountain regions than in lowlands, influenced more by socio-demographic factors than by wolf recolonisation. Localised shifts in acceptance appeared in clusters across the landscape, linked more to respondents' spatial proximity than to exposure to wolves or depredation events. Our findings highlight the importance of a nuanced approach to wildlife management that recognises areas of potential socioecological conflict as priorities for conservation, and of promoting societal discourse on community beliefs and values in landscapes of contested wolf presence.

Phenotype-dependent demography may result from heterogeneous selection pressures, which can be altered by temperature anomalies. Understanding which phenotypes are best adapted to different temperatures could clarify how populations may adapt to climatic changes and guide effective conservation measures. Animal markings are heritable phenotypic traits that show variation within populations, which suggest they may be adaptive. Variation in marking shape and size may influence thermoregulation and lead to differing individual responses to temperature anomalies. Therefore, markings may affect survival in variable and changing environments and be important for adapting to climate change. Using resighting data from 810 wild giraffes over 8 years, we estimated viability selection on spot patterns and whether it was affected by temperature anomalies. Calves with smaller lobate and adult males with smaller lobate or larger polygonal spots survived better. Additionally, calves and adult males with larger (vs. smaller) spots that experienced anomalously low (vs. high) temperatures survived better. Spot patterns had a smaller effect on adult females, with individuals of all spot types having lower survival in anomalously high temperatures. Synthesis and applications. Spot patterns influenced giraffe survival and their effects were altered by temperature anomalies. In calves, spot size may help with thermoregulation while spot shape may conceal them from predators. In adults, sex-specific selection pressures suggest sex differences in heat tolerance and trade-off with different functions. Conservation management maintaining variation in spot patterns by facilitating genetic exchange (e.g. through habitat connectivity) may help giraffes to adapt to climate change. Markings may have fitness consequences in other mammalian species and be important for population adaptation.

Habitat suitability models are commonly used to assess the potential distribution of large carnivores by identifying ecologically favourable areas. However, these models often overlook human dimensions, such as conflict and acceptance, which can lead to overestimation of species ranges and a mismatch between predicted and actual distributions. This limitation is particularly relevant for species like the snow leopard (Panthera uncia), which often inhabit areas shared with humans, where conflicts, especially livestock depredation, can trigger retaliatory killings. In this study, we integrated ecological suitability (ES) with human acceptance (HA) to identify socio-ecologically suitable habitats for snow leopards in northern Bhutan. HA was assessed through a questionnaire survey of 1046 households, capturing socio-demographic and geographic factors. We created a georeferenced map of HA and combined it with an existing ES map. Our analysis revealed that acceptance was higher among men and individuals aware of legal penalties for killing snow leopards but declined with increasing elevation and livestock losses. Agricultural farmers exhibited greater acceptance than livestock herders, Cordyceps mushroom collectors and those with other sources of livelihoods. The combined map revealed a strong overlap between socially and ecologically suitable areas, highlighting zones where both suitable ecological characteristics and community acceptance align. Although ES and HA generally aligned, our approach helped identify priority areas where targeted conservation interventions could reduce human-wildlife conflict. These findings underscore the value of incorporating human dimensions into habitat modelling. Such a holistic framework, combining ecological and social factors, can enhance conservation planning and promote human-carnivore coexistence, ultimately supporting the long-term persistence of snow leopard populations in northern Bhutan.Read the free for this article on the Journal blog.

Lack's seminal work on bird clutch sizes has spurred expansive research on reproductive trade-offs, especially focusing on offspring quantity-quality trade-offs and the potential fitness consequences for the parents. The environment is a critical driver of the expression of individual reproductive traits, influencing them through plastic responses. However, the plasticity of reproductive trade-offs themselves across environments has seldom been studied, and these studies were often limited to experimental approaches and dichotomous environments. Using 58 years of detailed data from a great tit population, we employ the recently developed "covariance reaction norm" (CRN) model to explore how continuous environmental variation influences the shape of reproductive trade-offs among individuals. Our analysis reveals that the correlation potentially indicative of the offspring quantity-quality trade-off is predominantly stable across years, with minimal variation linked to ecological harshness during the breeding season. However, the CRN also demonstrated that, despite some uncertainty associated with the results, the correlation between offspring mass and future offspring recruitment was positive, but only under harsh environmental conditions, suggesting that producing larger offspring provides fitness benefits when breeding conditions are suboptimal, which may reflect the importance of size for early-life competition. Altogether, this work highlights that there is temporal variation in some of the phenotypic correlations. This is a consequence of variation in offspring investment across breeding seasons, which is mostly driven by environmental conditions. Our study shows the benefits of exploring old ecological questions in the light of new statistical methods, highlighting the importance of understanding how environmental variation shapes the expression of life history trade-offs and the evolution of plasticity in reproductive strategies.

Reproductive success is a key determinant of individual fitness. Consequently, there has been a strong focus on identifying factors influencing mating success. Variation in mating success is often shaped by a combination of intrinsic traits, such as body size, age or behaviour, and extrinsic factors, such as environmental conditions and resource availability. Intense intrasexual competition, which commonly occurs in explosive-breeding amphibians due to male-skewed sex ratios at the breeding site, amplifies the importance of these traits and behaviours in gaining a competitive advantage. Using a 41-year individual-based capture–mark–recapture data set, we analysed the reproductive patterns of an alpine population of the common toad, Bufo bufo. We tested the effects of body size and age on the probability of a male being paired and being displaced while accounting for density and sex ratio. Additionally, we investigated the potential presence of intermittent breeding and whether paired males have different survival rates compared to unpaired males. Male body size was positively correlated with the probability of being paired and negatively correlated with displacement, showing that larger males are more successful in securing mates. Age negatively affected mating probability, suggesting reproductive senescence. Increasing male bias in the sex ratio led to a lower probability for males to be paired, while the population density did not affect the probability of pairing success. We found no evidence for intermittent breeding in males or differences in survival between paired and unpaired males. However, first-time breeding males had reduced survival compared to older males. This long-term field study provides one of the rare long-term perspectives on reproductive dynamics in amphibians, confirming experimental findings and highlighting the crucial role of body size and age as key life-history traits that determine reproductive success in this taxon.

Genetic diversity and structure are rarely assessed in populations established through conservation translocation. Here, we analysed the genetic structure and diversity of populations of an endangered pond-breeding amphibian, the common midwife toad, Alytes obstetricans, by comparing translocated populations against two types of populations: (i) populations which have recently colonized newly created ponds and (ii) natural populations which have been known to be present for a long time. Bottleneck events and dispersal patterns were analysed to describe the outcome of the translocations. In addition, we simulated trajectories of genetic diversity (He) of populations over time. The genetic diversity of natural colonized and translocated populations was similar to that of natural populations. However, there were signatures of genetic bottleneck events in three colonized populations and in a natural population. Simulations of genetic diversity over time showed that number and frequency of dispersers and population size are important parameters determining genetic diversity in the populations in the future. We conclude that, translocated, natural and colonized populations are genetically comparable, indicating that translocations can serve as an effective tool in conservation efforts. However, constructing ponds and waiting for natural colonization is also known to work well.

Responses of natural populations to climate change are driven by how multiple climatic and biotic factors affect survival and reproduction, and ultimately shape population dynamics. Yet, despite substantial progress in synthesizing the sensitivity of populations to climatic variation, comparative studies still overlook such complex interactions among drivers that generate variation in population-level metrics. Here, we use a common framework to synthesize how the joint effects of climate and biotic drivers on different vital rates impact population change, using unique long-term data from 41 species, ranging from trees to primates. We show that simultaneous effects of multiple climatic drivers exacerbate population responses to climate change, especially for fast-lived species. However, accounting for density feedbacks under climate variation buffers the effects of climate change on population dynamics. In all species considered in our analyses, such interactions between climate and density had starkly different effects depending on the age, size, or life-cycle stage of individuals, regardless of the life history of species. Our work provides the first general framework to assess how covarying effects of climate and density across a wide range of population models can impact populations of plants and animals under climate change.

Translocations are a commonly used method in conservation practice and are often viewed as successful if a new population persists. However, there is a need to understand the genetic and demographic consequences of translocations to better understand why some translocations are successful while others fail. Using a spatially structured population of the natterjack toad (Epidalea calamita) from northern Switzerland, this study presents a genetic and phenotypic analysis of natural and translocated populations. We used microsatellites to analyze the genetic structure (genetic diversity, differentiation, gene flow, and bottlenecks) of 102 individuals from 5 natural/donor and 64 individuals from 3 translocated populations. To study phenotypes (i.e., life history traits and behavior) of individuals, we used a common garden experiment. All genetic diversity metrics showed no genetic differences between natural and translocated populations with more genetic variation within than among populations. Although we found signatures of bottlenecks, they did not seem to have significantly affected genetic diversity of both natural and translocated populations. Life history traits of tadpoles (i.e., mass at metamorphosis, days to metamorphosis, survival) and behavior (tadpole activity) were similar in natural and translocated individuals. We conclude that the translocated populations have a genetic structure and life history traits similar to the donor and natural populations in the area. Overall, the results suggest that the approach used in this translocation program (individuals were taken from multiple donor populations and multiple females within populations and a large number of tadpoles in late developmental stages was released) was successful because it led to the establishment of populations that had levels of genetic diversity comparable to natural populations.

In most ecosystems, the increasingly strong effects of climate change on biodiversity co-occur with other anthropogenic pressures, most importantly land-use change. However, many long-term demographic studies focus on populations monitored in protected areas, and our understanding of how climate change will affect population persistence under anthropogenic land use is still limited. To fill this knowledge gap, we assessed the consequences of co-occurring land-use and climate change on vital rates and population dynamics of a fire-adapted Mediterranean carnivorous subshrub, the dewy pine (Drosophyllum lusitanicum). We used 7 years of individual data on 4753 plants monitored in three natural heathland sites that experience primarily fire as a disturbance, and five anthropogenic sites, where fires have been replaced by persistent disturbances from browsing or mechanical vegetation removal as a consequence of land-use change. All sites are projected to experience increasingly hotter summers and drier falls and winters. We used generalised additive models to model non-linear responses of survival, growth and reproduction to rainfall, temperature, size, density and time since fire in anthropogenic and natural dewy-pine populations. We then projected population dynamics under climate-change scenarios using an individual-based model. Our findings reveal that vital rates respond differently to climate change in anthropogenic compared to natural habitats. While extinction risks did not change under climate change in natural habitats, future higher summer temperatures decreased survival and led to population declines and higher extinction probabilities in anthropogenic habitats. Synthesis. Our results highlight the possible dramatic effects of climate change on populations largely confined to chronically disturbed, anthropogenic habitats and provide a foundation for devising relevant management strategies aiming towards the protection of species in human-disturbed habitats of the Mediterranean habitat. Overall, our findings emphasize the need for more long-term studies in managed landscapes.

Rodent body size often exhibits a diversity of temperature–size patterns among and within species, which might be caused by differential thermoneutral zones (TNZs) and experienced extents of warming. Here, we test this hypothesis in populations of Apodemus agrarius. To study how body size varies across space, we analysed data on body size and temperature (specifically, annual mean minimum temperature) from nine sites spanning 1150 m of elevation. Using indirect calorimetry, we also measured the resting metabolic rate at different temperatures to infer the population-specific TNZ. To study how body size changes over time (2013–2020), we analysed body-size data of southern and northern populations from warm and cold sites, respectively. With increasing temperatures across space, body size increased and the TNZ narrowed. Moreover, during the eight years, temperature remained stable at the warm site but rose at the cold site. As a result, body size increased in the population at the cold site but remained stable in the population at the warm site. Finally, the rate of change in body size per 1 °C change in temperature was larger along the temporal than the spatial temperature gradient. Together, these results support our hypothesis that, among rodent populations, differential changes in body size can be caused by site-specific habitat warming and the population-specific TNZ. A population with a narrow TNZ can be restricted in its body-size response to habitat warming.

Survival, reproduction, and movement are the key demographic parameters that drive population dynamics. Factors affecting these demographic parameters in large, long‐lived, extinction‐threatened mammals are diverse and may differentially affect subpopulations in disparate parts of an ecosystem. We conducted annual photographic surveys to uniquely identify 1,520 giraffes at 4 subpopulations around the Serengeti Ecosystem in Tanzania to estimate demographic parameters of age‐ and sex‐specific survival probabilities, reproduction, population densities, group sizes, and long‐distance movements. In the Seronera (central) subpopulation, we combined 15 years of data from 3 independent survey schemes, developed a Bayesian hidden Markov model to estimate demographic parameters, and conducted a retrospective population analysis to elucidate the demographic drivers of temporal changes in population growth rate. We collected data over 4–5 years for 3 other subpopulations, and used frequentist methods to estimate demographic parameters. We compared our results with historical estimates from the 1970s and 2000s to examine long‐term population trends and demographic drivers. We found significant differences in adult and subadult survival probabilities among subpopulations, with lower adult survival associated with declining subpopulations. Retrospective population analysis for the Seronera subpopulation reiterated that adult survival is a critical demographic driver of population dynamics for giraffes. The 2 subpopulations adjacent to the protected area boundary declined over 48 years, whereas the Seronera subpopulation stabilized since 2008. Only one individual moved between subpopulations, providing evidence for subpopulation insularity and potential genetic structuring of the overall population. These factors underscore the need for subpopulation‐specific conservation strategies aimed at raising adult survival within the western and northeastern parts of the Serengeti Ecosystem. Community‐based conservation efforts adjacent to protected areas have been effective in raising adult survival and density elsewhere. Our findings highlight the importance of understanding subpopulation dynamics and their demographic drivers for evidence‐based conservation and management to recover endangered giraffe populations.

Most mammals, including humans, exhibit even or slightly male-biased birth sex ratios (BSRs) and female-biased adult sex ratios (ASRs) much later in life due to higher male mortality rates. The group-living primates of Madagascar are unusual in this respect because they lack female-biased ASRs, but it is unknown whether this is the result of skewed BSRs or sex-specific disappearance patterns. Using long-term demographic data from wild red-fronted lemurs ( Eulemur rufifrons ), we analysed their sex ratio dynamics across the lifespan. We assessed BSR via prenatal sex determination using maternal faecal oestrogen metabolite measurements during late pregnancy, confirming a visually determined equal sex ratio three months after birth, and indicating no early sex-specific mortality. Demographic analyses additionally disclosed higher female disappearance within the first 8 years of age, likely associated with reproductive effort early in life. Thereby, adult male survival had the greatest positive effect on the ASR. Our study offers a rare perspective on the dynamics of age- and sex-specific disappearance in a wild primate population, whose sex-reversed patterns may also contribute to a more general understanding of the mechanisms generating sex-biased mortality.

Alpine species are severely affected by climate change, with elevational range shifts being one key response of mountain species to the rapidly warming environment. The Alpine marmot (Marmota marmota) is suggested to be particularly susceptible to ongoing warming. However, it is largely unknown how climate change affected the Alpine marmot distribution in recent decades. This study examines the elevational changes in Alpine marmot distribution over the past 40 years in a Central Alps Mountain valley. Based on historical occurrence data of the year 1982, we resurveyed the marmot occurrences in the year 2022. We analysed potential distributional changes over time by fitting dynamic site-occupancy models to detect occupancy patterns, as well as marmot colonisations and site abandonments ('local extinctions' at a site) along the elevational gradient, whilst accounting for imperfect detection. Contrary to expectations, we found no evidence of upward colonisation at higher elevations or an upward shift of the lower range margin in our study, suggesting that marmots are not climate-limited at lower elevations in the investigated valley, and other factors than climate might constrain their higher elevation colonisation. Nevertheless, the marmot's elevational optimum shifted upwards by +86 m. Our results indicate that the most favourable conditions for marmots have slightly shifted higher due to warming. To better understand potential habitat contractions driven by climate change, further large-scale studies focusing on the lower range margins in warmer Alpine regions are necessary. Recognising distribution changes of species vulnerable to climate change is crucial to evaluate local extinction risks and for conserving biodiversity.

Aim: Groundwater ecosystems harbour a unique biodiversity, but remain poorly studied, mainly due to difficulties in accessibility and imperfect species detection. Consequently, knowledge on the state and change of groundwater biodiversity remains highly deficient. In the context of global warming and excessive groundwater extraction, understanding groundwater from an ecosystem-perspective, including organism diversity and distribution, is essential. This study presents the largest ever systematic assessment of groundwater amphipods, which are a key component of European groundwater biodiversity. Location: Switzerland (41,285 km2), including data from 906 sampling sites. Taxon: Groundwater amphipods, genera Niphargus and Crangonyx (Crustacea, Amphipoda). Methods: We applied a highly standardized citizen science approach to collect repeated groundwater fauna samples in collaboration with municipal drinking water providers. Using detection-nondetection data of the genetically identified groundwater amphipod species, we assessed the overall species diversity of both rare and common species. The distribution of commonly found species was predicted using multispecies occupancy modelling. Results: We retrieved 3882 samples from 906 sites, yielding 2350 groundwater amphipod individuals. We identified a remarkable species diversity, comprising few commonly and many rarely found species. Considering commonly found species, we identified distinct distribution ranges, low local species richness and a predominance of negative co-occurrences. In contrast, a major portion of species were found rarely (generally at just one or two sites each), distributed uniformly throughout the study area and unrelated to common species' recognized hotspots. Many of these rarely found species are not yet formally described. Main Conclusions: Our results give robust emphasis on the rare occurrence and narrow distribution of many groundwater dwellers. Our systematic and standardized sampling data of groundwater amphipods suggest that rarity is particularly prominent and inherent to groundwater organisms. We emphasize the need of systematic data to integrate rare groundwater species in biodiversity assessments, especially in times of global change.

Density dependent carry-over effects from one life history stage to another can affect the dynamics of populations. Here we study such carry-over effects from the tadpole to the postmetamorphic juvenile stage in an endangered amphibian, the natterjack toad (Epidalea calamita). We raised tadpoles in outdoor aquatic mesocosms at four densities and assessed juvenile performance after metamorphosis in terrestrial mesocosms. High larval density reduced mass at metamorphosis by 50 % and doubled the length of the larval period. Survival was reduced at the high densities. Larger metamorphs had higher survival in terrestrial mesocosms and remained larger than cohort members at the end of the 30-day experiment. Because juvenile survival drives amphibian population dynamics, density-dependent carry-over effects to the juvenile stage are likely to affect population viability. We discuss the implications of the results for amphibian conservation practice, both pond construction programs and surveys of amphibian populations.

While climate change has been shown to impact several life-history traits of wild-living animal populations, little is known about its effects on dispersal and connectivity. Here, we capitalize on the highly variable flooding regime of the Okavango Delta to investigate the impacts of changing environmental conditions on the dispersal and connectivity of the endangered African wild dog (Lycaon pictus). Based on remote sensed flood extents observed over 20 years, we derive two extreme flood scenarios: a minimum and a maximum flood extent, representative of very dry and very wet environmental periods. These conditions are akin to those anticipated under increased climatic variability, as it is expected under climate change. Using a movement model parameterized with GPS data from dispersing individuals, we simulate 12,000 individual dispersal trajectories across the ecosystem under both scenarios and investigate patterns of connectivity. Across the entire ecosystem, surface water coverage during maximum flood extent reduces dispersal success (i.e., the propensity of individuals to disperse between adjacent subpopulations) by 12% and increases dispersal durations by 17%. Locally, however, dispersal success diminishes by as much as 78%. Depending on the flood extent, alternative dispersal corridors emerge, some of which in the immediate vicinity of human-dominated landscapes. Notably, under maximum flood extent, the number of dispersing trajectories moving into human-dominated landscapes decreases by 41% at the Okavango Delta's inflow, but increases by 126% at the Delta's distal end. This may drive the amplification of human-wildlife conflict. While predicting the impacts of climate change on environmental conditions on the ground remains challenging, our results highlight that environmental change may have significant consequences for dispersal patterns and connectivity, and ultimately, population viability. Acknowledging and anticipating such impacts will be key to effective conservation strategies and to preserve vital dispersal corridors in light of climate change and other human-related landscape alterations.

AimVariation in community composition along environmental gradients provides crucial information for identifying zones where species turnover is rapid and to ascertain whether compositional changes occur gradually or rather abruptly. We examined changes in bird community composition along three bioclimatic transects in Australia to test whether drivers of species turnover are consistent, rather than spatially contingent, across biologically contrasting ecosystems. We also detected potential transition zones associated with environmental thresholds and determined whether certain abiotic conditions promote a higher rate of community compositional turnover.LocationMainland Australia.TaxonTerrestrial birds.MethodsWe applied multivariate community analysis, generalised dissimilarity modelling (GDM) and threshold indicator taxa analysis (TITAN).ResultsWe observed that environmental variables are better predictors of community composition than spatial distance, which indicates that species sorting, rather than dispersal, plays a key role in structuring Australian avian communities. Annual precipitation constitutes a key driver of species turnover regardless of the analysed transect. The most humid landscapes and those with a higher tree canopy show lower spatial heterogeneity in community composition compared to those with less benign environmental conditions (e.g., dryer environments). TITAN detected significant transition points and supported the results obtained using GDM, which suggests that bird composition change along the gradients is not monotonic.Main ConclusionsOur results suggest that avian beta diversity increases with increasing environmental harshness, presumably through changes in the relative importance of stochastic versus deterministic processes. The obtained findings show that open forests and woodlands are extremely important ecosystems on this continent and deserve special attention in terms of conservation due to their vulnerability to global change. Lastly, this study exemplifies the value of combining community- and taxon-based analyses to identify and interpret community thresholds, which can serve to pinpoint targets for preserving biodiversity.

1. Vital rates of individuals in natural populations, such as survival and breeding probabilities, can be affected by abiotic (i.e. environmental conditions), biotic (e.g. population density), and individual factors (i.e. individual traits). Many studies often consider the direct effects of one or two of these sources of vital-rate variations, but taking them all into account might reveal important relationships, including indirect trait-mediated effects. 2. We estimated survival and breeding probability in a high-elevation population of an ectotherm species while accounting for the effects of these three types of factors: abiotic (in the form of the length of the active season and temperature at emergence from hibernation), biotic (population size), and individual factors (body size). We expect this population living at the extreme range of the species’ distribution to be affected mainly by abiotic factors, given the harsh environmental conditions experienced. 3. Using 28 years of capture-mark-recapture data and individual body size measurements on an alpine population of the common toad (Bufo bufo), we estimated male survival probability with a Bayesian Cormack-Jolly-Seber model, female survival and female breeding probability with a Bayesian multistate model, and we estimated sex-specific growth curves. 4. Our findings reveal significant negative impacts of all three factor types on survival. Moreover, breeding probability exhibited a Markovian pattern, notably breeding biennially, with temperature at hibernation emergence, an abiotic factor, affecting the probability of skipping a breeding season, and population density, a biotic factor, influencing the resuming of breeding. We also found indication for the presence of indirect effects, with both abiotic and biotic factors potentially affecting asymptotic growth, and thus by extension, survival through changes in body size. 5. Our study reveals interesting and novel results for a population in a context seldom studied. It also highlights the complexity of factors affecting vital rates directly and indirectly, as well as the importance of long-term studies, especially in understudied habitats and taxa.