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Interactions between density and environmental conditions have important effects on vital rates and consequently on population dynamics and can take complex pathways in species whose demography is strongly influenced by social context, such as the African lion, Panthera leo. In populations of such species, the response of vital rates to density can vary depending on the social structure (e.g. effects of group size or composition). However, studies assessing density dependence in populations of lions and other social species have seldom considered the effects of multiple socially explicit measures of density, and-more particularly for lions-of nomadic males. Additionally, vital-rate responses to interactions between the environment and various measures of density remain largely uninvestigated. To fill these knowledge gaps, we aimed to understand how a socially and spatially explicit consideration of density (i.e. at the local scale) and its interaction with environmental seasonality affect vital rates of lions in the Serengeti National Park, Tanzania. We used a Bayesian multistate capture-recapture model and Bayesian generalized linear mixed models to estimate lion stage-specific survival and between-stage transition rates, as well as reproduction probability and recruitment, while testing for season-specific effects of density measures at the group and home-range levels. We found evidence for several such effects. For example, resident-male survival increased more strongly with coalition size in the dry season compared with the wet season, and adult-female abundance affected subadult survival negatively in the wet season, but positively in the dry season. Additionally, while our models showed no effect of nomadic males on adult-female survival, they revealed strong effects of nomads on key processes such as reproduction and takeover dynamics. Therefore, our results highlight the importance of accounting for seasonality and social context when assessing the effects of density on vital rates of Serengeti lions and of social species more generally., (© 2024 The Author(s). Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Life history trade-offs are one of the central tenets of evolutionary demography. Trade-offs, depicting negative covariances between individuals' life history traits, can arise from genetic constraints, or from a finite amount of resources that each individual has to allocate in a zero-sum game between somatic and reproductive functions. While theory predicts that trade-offs are ubiquitous, empirical studies have often failed to detect such negative covariances in wild populations. One way to improve the detection of trade-offs is by accounting for the environmental context, as trade-off expression may depend on environmental conditions. However, current methodologies usually search for fixed covariances between traits, thereby ignoring their context dependence. Here, we present a hierarchical multivariate 'covariance reaction norm' model, adapted from Martin (2023), to help detect context dependence in the expression of life-history trade-offs using demographic data. The method allows continuous variation in the phenotypic correlation between traits. We validate the model on simulated data for both intraindividual and intergenerational trade-offs. We then apply it to empirical datasets of yellow-bellied marmots (Marmota flaviventer) and Soay sheep (Ovis aries) as a proof-of-concept showing that new insights can be gained by applying our methodology, such as detecting trade-offs only in specific environments. We discuss its potential for application to many of the existing long-term demographic datasets and how it could improve our understanding of trade-off expression in particular, and life history theory in general., (© 2024 The Author(s). Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Early warning signals (EWS) are phenomenological tools that have been proposed as predictors of the collapse of biological systems. Although a growing body of work has shown the utility of EWS based on either statistics derived from abundance data or shifts in phenotypic traits such as body size, so far this work has largely focused on single species populations. However, to predict reliably the future state of ecological systems, which inherently could consist of multiple species, understanding how reliable such signals are in a community context is critical. Here, reconciling quantitative trait evolution and Lotka-Volterra equations, which allow us to track both abundance and mean traits, we simulate the collapse of populations embedded in mutualistic and multi-trophic predator-prey communities. Using these simulations and warning signals derived from both population- and community-level data, we showed the utility of abundance-based EWS, as well as metrics derived from stability-landscape theory (e.g. width and depth of the basin of attraction), were fundamentally linked. Thus, the depth and width of such stability-landscape curves could be used to identify which species should exhibit the strongest EWS of collapse. The probability a species displays both trait and abundance-based EWS was dependent on its position in a community, with some species able to act as indicator species. In addition, our results also demonstrated that in general trait-based EWS were less reliable in comparison with abundance-based EWS in forecasting species collapses in our simulated communities. Furthermore, community-level abundance-based EWS were fairly reliable in comparison with their species-level counterparts in forecasting species-level collapses. Our study suggests a holistic framework that combines abundance-based EWS and metrics derived from stability-landscape theory that may help in forecasting species loss in a community context., (© 2022 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

The ability of dispersing individuals to adjust their behaviour to changing conditions is instrumental in overcoming challenges and reducing dispersal costs, consequently increasing overall dispersal success. Understanding how dispersers' behaviour and physiology change during the dispersal process, and how they differ from resident individuals, can shed light on the mechanisms by which dispersers increase survival and maximise reproduction. By analysing individual behaviour and concentrations of faecal glucocorticoid metabolites (fGCM), a stress-associated biomarker, we sought to identify the proximate causes behind differences in survival and reproduction between dispersing and resident meerkats Suricata suricatta. We used data collected on 67 dispersing and 108 resident females to investigate (a) which individual, social and environmental factors are correlated to foraging and vigilance, and whether the role of such factors differs among dispersal phases, and between dispersers and residents; (b) how time allocated to either foraging or vigilance correlated to survival in dispersers and residents and (c) the link between aggression and change in fGCM concentration, and their relationship with reproductive rates in dispersing groups and resident groups with either long-established or newly established dominant females. Time allocated to foraging increased across dispersal phases, whereas time allocated to vigilance decreased. Time allocated to foraging and vigilance correlated positively and negatively, respectively, with dispersers' group size. We did not find a group size effect for residents. High proportions of time allocated to foraging correlated with high survival, and more so in dispersers, suggesting that maintaining good physical condition may reduce mortality during dispersal. Furthermore, while subordinate individuals rarely reproduced in resident groups, the conception rate of subordinates in newly formed dispersing groups was equal to that of their dominant individuals. Mirroring conception rates, in resident groups, fGCM concentrations were lower in subordinates than in dominants, whereas in disperser groups, fGCM concentrations did not differ between subordinates and dominants. Our results, which highlight the relationship between behavioural and physiological factors and demographic rates, provide insights into some of the mechanisms that individuals of a cooperative species can use to increase overall dispersal success., (© 2021 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Approximately 25% of mammals are currently threatened with extinction, a risk that is amplified under climate change. Species persistence under climate change is determined by the combined effects of climatic factors on multiple demographic rates (survival, development and reproduction), and hence, population dynamics. Thus, to quantify which species and regions on Earth are most vulnerable to climate-driven extinction, a global understanding of how different demographic rates respond to climate is urgently needed. Here, we perform a systematic review of literature on demographic responses to climate, focusing on terrestrial mammals, for which extensive demographic data are available. To assess the full spectrum of responses, we synthesize information from studies that quantitatively link climate to multiple demographic rates. We find only 106 such studies, corresponding to 87 mammal species. These 87 species constitute <1% of all terrestrial mammals. Our synthesis reveals a strong mismatch between the locations of demographic studies and the regions and taxa currently recognized as most vulnerable to climate change. Surprisingly, for most mammals and regions sensitive to climate change, holistic demographic responses to climate remain unknown. At the same time, we reveal that filling this knowledge gap is critical as the effects of climate change will operate via complex demographic mechanisms: a vast majority of mammal populations display projected increases in some demographic rates but declines in others, often depending on the specific environmental context, complicating simple projections of population fates. Assessments of population viability under climate change are in critical need to gather data that account for multiple demographic responses, and coordinated actions to assess demography holistically should be prioritized for mammals and other taxa., (© 2021 British Ecological Society.)

Experimental laboratory evidence suggests that animals with disrupted social systems express weakened relationship strengths and have more exclusive social associations, and that these changes have functional consequences. A key question is whether anthropogenic pressures have a similar impact on the social structure of wild animal communities. We addressed this question by constructing a social network from 6 years of systematically collected photographic capture-recapture data spanning 1,139 individual adult female Masai giraffes inhabiting a large, unfenced, heterogeneous landscape in northern Tanzania. We then used the social network to identify distinct social communities, and tested whether social or anthropogenic and other environmental factors predicted differences in social structure among these communities. We reveal that giraffes have a multilevel social structure. Local preferences in associations among individuals scale up to a number of distinct, but spatially overlapping, social communities, that can be viewed as a large interconnected metapopulation. We then find that communities that are closer to traditional compounds of Indigenous Masai people express weaker relationship strengths and the giraffes in these communities are more exclusive in their associations. The patterns we characterize in response to proximity to humans reflect the predictions of disrupted social systems. Near bomas, fuelwood cutting can reduce food resources, and groups of giraffes are more likely to encounter livestock and humans on foot, thus disrupting the social associations among group members. Our results suggest that human presence could potentially be playing an important role in determining the conservation future of this megaherbivore., (© 2020 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of British Ecological Society.)

Reliable estimates of birth, death, emigration and immigration rates are fundamental to understanding and predicting the dynamics of wild populations and, consequently, inform appropriate management actions. However, when individuals disappear from a focal population, inference on their fate is often challenging. Here we used 30 years of individual-based mark-recapture data from a population of free-ranging African wild dogs Lycaon pictus in Botswana and a suite of individual, social and environmental predictors to investigate factors affecting the decision to emigrate from the pack. We subsequently used this information to assign an emigration probability to those individuals that were no longer sighted within their pack (i.e. missing individuals). Natal dispersal (i.e. emigration from the natal pack) showed seasonal patterns with female dispersal peaking prior to the mating season and male dispersal peaking at the beginning of the wet season. For both sexes, natal dispersal rate increased in the absence of unrelated individuals of the opposite sex in the pack. Male natal dispersal decreased with increasing number of pups in the pack and increased in larger packs. Female natal dispersal decreased with increasing number of pups in larger packs, but increased with increasing number of pups in smaller packs. Individuals of both sexes were less likely to exhibit secondary dispersal (i.e. emigration from a pack other than the natal pack) if they were dominant and if many pups were present in the pack. Our models predicted that 18% and 25% of missing females and males, respectively, had likely dispersed from the natal pack, rather than having died. A misclassification of this order of magnitude between dispersal and mortality can have far-reaching consequences in the evaluation and prediction of population dynamics and persistence, and potentially mislead conservation actions. Our study showed that the decision to disperse is context-dependent and that the effect of individual, social and environmental predictors differs between males and females and between natal and secondary dispersal related to different direct and indirect fitness consequences. Furthermore, we demonstrated how a thorough understanding of the proximate causes of dispersal can be used to assign a dispersal probability to missing individuals. Knowledge of causes of dispersal can then be used within an integrated framework to more reliably estimate mortality rates., (© 2020 British Ecological Society.)

Environmental change can impact the stability of ecological systems and cause rapid declines in populations. Abundance-based early warning signals have been shown to precede such declines, but detection prior to wild population collapses has had limited success, leading to the development of warning signals based on shifts in distribution of fitness-related traits such as body size. The dynamics of population abundances and traits in response to external environmental perturbations are controlled by a range of underlying factors such as reproductive rate, genetic variation and plasticity. However, it remains unknown how such ecological and evolutionary factors affect the stability landscape of populations and the detectability of abundance and trait-based early warning signals. Here, we apply a trait-based demographic approach and investigate both trait and population dynamics in response to gradual and increasing changes in the environment. We explore a range of ecological and evolutionary constraints under which stability of a population may be affected. We show both analytically and with simulations that strength of abundance- and trait-based warning signals are affected by ecological and evolutionary factors. Finally, we show that combining trait- and abundance-based information improves our ability to predict population declines. Our study suggests that the inclusion of trait dynamic information alongside generic warning signals should provide more accurate forecasts of the future state of biological systems., (© 2019 The Authors. Journal of Animal Ecology © 2019 British Ecological Society.)

Population dynamics are the result of an interplay between extrinsic and intrinsic environmental drivers. Predicting the effects of environmental change on wildlife populations therefore requires a thorough understanding of the mechanisms through which different environmental drivers interact to generate changes in population size and structure. In this study, we disentangled the roles of temperature, food availability and population density in shaping short- and long-term population dynamics of the African striped mouse, a small rodent inhabiting a semidesert with high intra- and interannual variation in environmental conditions. We parameterized a female-only stage-structured matrix population model with vital rates depending on temperature, food availability and population density, using monthly mark-recapture data from 1609 mice trapped over 9 years (2005-2014). We then applied perturbation analyses to determine relative strengths and demographic pathways of these drivers in affecting population dynamics. Furthermore, we used stochastic population projections to gain insights into how three different climate change scenarios might affect size, structure and persistence of this population. We identified food availability, acting through reproduction, as the main driver of changes in both short- and long-term population dynamics. This mechanism was mediated by strong density feedbacks, which stabilized the population after high peaks and allowed it to recover from detrimental crashes. Density dependence thus buffered the population against environmental change, and even adverse climate change scenarios were predicted to have little effect on population persistence (extinction risk over 100 years <5%) despite leading to overall lower abundances. Explicitly linking environment-demography relationships to population dynamics allowed us to accurately capture past population dynamics. It further enabled establishing the roles and relative importances of extrinsic and intrinsic environmental drivers, and we conclude that doing this is essential when investigating impacts of climate change on wildlife populations., (© 2018 The Authors. Journal of Animal Ecology © 2018 British Ecological Society.)

Dispersal is a key process governing the dynamics of socially and spatially structured populations and involves three distinct stages: emigration, transience and settlement. At each stage, individuals have to make movement decisions, which are influenced by social, environmental and individual factors. Yet, a comprehensive understanding of the drivers that influence such decisions is still lacking, particularly for the transient stage during which free-living individuals are inherently difficult to follow. Social circumstances such as the likelihood of encountering conspecifics can be expected to strongly affects decision-making during dispersal, particularly in territorial species where encounters with resident conspecifics are antagonistic. Here, we analysed the movement trajectories of 47 dispersing coalitions of Kalahari meerkats Suricata suricatta through a landscape occupied by constantly monitored resident groups, while simultaneously taking into account environmental and individual characteristics. We used GPS locations collected on resident groups to create a georeferenced social landscape representing the likelihood of encountering resident groups. We used a step-selection function to infer the effect of social, environmental and individual covariates on habitat selection during dispersal. Finally, we created a temporal mismatch between the social landscape and the dispersal event of interest to identify the temporal scale at which dispersers perceive the social landscape. Including information about the social landscape considerably improved our representation of the dispersal trajectory compared to analyses that only accounted for environmental variables. The latter were only marginally selected or avoided by dispersers. Before leaving their natal territory, dispersers selected areas frequently used by their natal group. In contrast, after leaving their natal territory, they selectively used areas where they were less likely to encounter unrelated groups. This pattern was particularly marked in larger dispersing coalitions and when unrelated males were part of the dispersing coalition. Our results suggest that, in socially and spatially structured species, dispersers gather and process social information during dispersal, and that reducing risk of aggression from unrelated resident groups outweighs benefits derived from conspecific attraction. Finally, our work underlines the intimate link between the social structure of a population and dispersal, which affect each other reciprocally., (© 2018 The Authors. Journal of Animal Ecology © 2018 British Ecological Society.)

Large-scale fluctuations in abundance are a common feature of small mammal populations and have been the subject of extensive research. These demographic fluctuations are often associated with concurrent changes in the average body mass of individuals, sometimes referred to as the 'Chitty effect'. Despite the long-standing recognition of this phenomenon, an empirical investigation of the underlying coupled dynamics of body mass and population growth has been lacking. Using long-term life-history data combined with a trait-based demographic approach, we examined the relationship between body mass and demography in a small mammal population that exhibits non-cyclic, large-scale fluctuations in abundance. We used data from the male segment of a 25-year study of the monogamous prairie vole, Microtus ochrogaster, in Illinois, USA. Specifically, we investigated how trait-demography relationships and trait distributions changed between different phases of population fluctuations, and the consequences of these changes for both trait and population dynamics. We observed phase-specific changes in male adult body mass distribution in this population of prairie voles. Our analyses revealed that these changes were driven by variation in ontogenetic growth, rather than selection acting on the trait. The resulting changes in body mass influenced most life-history processes, and these effects varied among phases of population fluctuation. However, these changes did not propagate to affect the population growth rate due to the small effect of body mass on vital rates, compared to the overall differences in vital rates between phases. The increase phase of the fluctuations was initiated by enhanced survival, particularly of juveniles and fecundity, whereas the decline phase was driven by an overall reduction in fecundity, survival and maturation rates. Our study provides empirical support, as well as a potential mechanism, underlying the observed trait changes accompanying population fluctuations. Body size dynamics and population fluctuations resulted from different life-history processes. Therefore, we conclude that body size dynamics in our population do not drive the observed population dynamics. This more in-depth understanding of different components of small mammal population fluctuations will help us to better identify the mechanistic drivers of this interesting phenomenon., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

Wind is an important climatic factor for flying animals as by affecting their locomotion, it can deeply impact their life-history characteristics. In the context of globally changing wind patterns, we investigated the mechanisms underlying recently reported increase in body mass of a population of wandering albatrosses (Diomedea exulans) with increasing wind speed over time. We built a foraging model detailing the effects of wind on movement statistics and ultimately on mass gained by the forager and mass lost by the incubating partner. We then simulated the body mass of incubating pairs under varying wind scenarios. We tracked the frequency at which critical mass leading to nest abandonment was reached to assess incubation success. We found that wandering albatrosses behave as time minimizers during incubation as mass gain was independent of any movement statistics but decreased with increasing mass at departure. Individuals forage until their energy requirements, which are determined by their body conditions, are fulfilled. This can come at the cost of their partner's condition as mass loss of the incubating partner depended on trip duration. This behaviour is consistent with strategies of long-lived species which favoured their own survival over their current reproductive attempt. In addition, wind speed increased ground speed which in turn reduced trip duration and males foraged further away than females at high ground speed. Contrasted against an independent data set, the simulation performed satisfactorily for males but less so for females under current wind conditions. The simulation predicted an increase in male body mass growth rate with increasing wind speed, whereas females' rate decreased. This trend may provide an explanation for the observed increase in mass of males but not of females. Conversely, the simulation predicted very few nest abandonments, which is in line with the high breeding success of this species and is contrary to the hypothesis that wind patterns impact incubation success by altering foraging movement., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

Demographic rates are shaped by the interaction of past and current environments that individuals in a population experience. Past environments shape individual states via selection and plasticity, and fitness-related traits (e.g. individual size) are commonly used in demographic analyses to represent the effect of past environments on demographic rates. We quantified how well the size of individuals captures the effects of a population's past and current environments on demographic rates in a well-studied experimental system of soil mites. We decomposed these interrelated sources of variation with a novel method of multiple regression that is useful for understanding nonlinear relationships between responses and multicollinear explanatory variables. We graphically present the results using area-proportional Venn diagrams. Our novel method was developed by combining existing methods and expanding upon them. We showed that the strength of size as a proxy for the past environment varied widely among vital rates. For instance, in this organism with an income breeding life history, the environment had more effect on reproduction than individual size, but with substantial overlap indicating that size encompassed some of the effects of the past environment on fecundity. This demonstrates that the strength of size as a proxy for the past environment can vary widely among life-history processes within a species, and this variation should be taken into consideration in trait-based demographic or individual-based approaches that focus on phenotypic traits as state variables. Furthermore, the strength of a proxy will depend on what state variable(s) and what demographic rate is being examined; that is, different measures of body size (e.g. length, volume, mass, fat stores) will be better or worse proxies for various life-history processes., (© 2016 The Authors. Journal of Animal Ecology © 2016 British Ecological Society.)

Social and environmental factors influence key life-history processes and population dynamics by affecting fitness-related phenotypic traits such as body mass. The role of body mass is particularly pronounced in cooperative breeders due to variation in social status and consequent variation in access to resources. Investigating the mechanisms underlying variation in body mass and its demographic consequences can help elucidate how social and environmental factors affect the dynamics of cooperatively breeding populations. In this study, we present an analysis of the effect of individual variation in body mass on the temporal dynamics of group size and structure of a cooperatively breeding mongoose, the Kalahari meerkat, Suricata suricatta. First, we investigate how body mass interacts with social (dominance status and number of helpers) and environmental (rainfall and season) factors to influence key life-history processes (survival, growth, emigration and reproduction) in female meerkats. Next, using an individual-based population model, we show that the models explicitly including individual variation in body mass predict group dynamics better than those ignoring this morphological trait. Body mass influences group dynamics mainly through its effects on helper emigration and dominant reproduction. Rainfall has a trait-mediated, destabilizing effect on group dynamics, whereas the number of helpers has a direct and stabilizing effect. Counteracting effects of number of helpers on different demographic rates, despite generating temporal fluctuations, stabilizes group dynamics in the long term. Our study demonstrates that social and environmental factors interact to produce individual variation in body mass and accounting for this variation helps to explain group dynamics in this cooperatively breeding population., (© 2014 The Authors. Journal of Animal Ecology © 2014 British Ecological Society.)

Resource availability plays a key role in driving variation in somatic growth and body condition, and the factors determining access to resources vary considerably across life stages. Parents and carers may exert important influences in early life, when individuals are nutritionally dependent, with abiotic environmental effects having stronger influences later in development as individuals forage independently. Most studies have measured specific factors influencing growth across development or have compared relative influences of different factors within specific life stages. Such studies may not capture whether early-life factors continue to have delayed effects at later stages, or whether social factors change when individuals become nutritionally independent and adults become competitors for, rather than providers of, food. Here, we examined variation in the influence of the abiotic, social and maternal environment on growth across life stages in a wild population of cooperatively breeding meerkats. Cooperatively breeding vertebrates are ideal for investigating environmental influences on growth. In addition to experiencing highly variable abiotic conditions, cooperative breeders are typified by heterogeneity both among breeders, with mothers varying in age and social status, and in the number of carers present. Recent rainfall had a consistently marked effect on growth across life stages, yet other seasonal terms only influenced growth during stages when individuals were growing fastest. Group size and maternal dominance status had positive effects on growth during the period of nutritional dependence on carers, but did not influence mass at emergence (at 1 month) or growth at independent stages (>4 months). Pups born to older mothers were lighter at 1 month of age and subsequently grew faster as subadults. Males grew faster than females during the juvenile and subadult stage only. Our findings demonstrate the complex ways in which the external environment influences development in a cooperative mammal. Individuals are most sensitive to social and maternal factors during the period of nutritional dependence on carers, whereas direct environmental effects are relatively more important later in development. Understanding the way in which environmental sensitivity varies across life stages is likely to be an important consideration in predicting trait responses to environmental change., (© 2013 The Authors. Journal of Animal Ecology published by John Wiley & Sons Ltd on behalf of the British Ecological Society.)

1. For social species, the link between individual behaviour and population dynamics is mediated by group-level demography. 2. Populations of obligate cooperative breeders are structured into social groups, which may be subject to inverse density dependence (Allee effects) that result from a dependence on conspecific helpers, but evidence for population-wide Allee effects is rare. 3. We use field data from a long-term study of cooperative meerkats (Suricata suricatta; Schreber, 1776) - a species for which local Allee effects are not reflected in population-level dynamics - to empirically model interannual group dynamics. 4. Using phenomenological population models, modified to incorporate environmental conditions and potential Allee effects, we first investigate overall patterns of group dynamics and find support only for conventional density dependence that increases after years of low rainfall. 5. To explain the observed patterns, we examine specific demographic rates and assess their contributions to overall group dynamics. Although per-capita meerkat mortality is subject to a component Allee effect, it contributes relatively little to observed variation in group dynamics, and other (conventionally density dependent) demographic rates - especially emigration - govern group dynamics. 6. Our findings highlight the need to consider demographic processes and density dependence in subpopulations before drawing conclusions about how behaviour affects population processes in socially complex systems., (© 2011 The Authors. Journal of Animal Ecology © 2011 British Ecological Society.)

1. The presence/absence of a species at a particular site is the simplest form of data that can be collected during ecological field studies. We used 13 years (1990-2002) of survey data to parameterize a stochastic patch occupancy model for a metapopulation of the yellow-bellied marmot in Colorado, and investigated the significance of particular patches and the influence of site quality, network characteristics and regional stochasticity on the metapopulation persistence. 2. Persistence of the yellow-bellied marmot metapopulation was strongly dependent on the high quality colony sites, and persistence probability was highly sensitive to small changes in the quality of these sites. 3. A relatively small number of colony sites was ultimately responsible for the regional persistence. However, lower quality satellite sites also made a significant contribution to long-term metapopulation persistence, especially when regional stochasticity was high. 4. The northern network of the marmot metapopulation was more stable compared to the southern network, and the persistence of the southern network depended heavily on the northern network. 5. Although complex models of metapopulation dynamics may provide a more accurate description of metapopulation dynamics, such models are data-intensive. Our study, one of the very few applications of stochastic patch occupancy models to a mammalian species, suggests that stochastic patch occupancy models can provide important insights into metapopulation dynamics using data that are easy to collect.