Behavioural responses to human‐induced environmental change

The initial response of individuals to human‐induced environmental change is often behavioural. This can improve the performance of individuals under sudden, large‐scale perturbations and maintain viable populations. The response can also give additional time for genetic changes to arise and, hence, facilitate adaptation to new conditions. On the other hand, maladaptive responses, which reduce individual fitness, may occur when individuals encounter conditions that the population has not experienced during its evolutionary history, which can decrease population viability. A growing number of studies find human disturbances to induce behavioural responses, both directly and by altering factors that influence fitness. Common causes of behavioural responses are changes in the transmission of information, the concentration of endocrine disrupters, the availability of resources, the possibility of dispersal, and the abundance of interacting species. Frequent responses are alterations in habitat choice, movements, foraging, social behaviour and reproductive behaviour. Behavioural responses depend on the genetically determined reaction norm of the individuals, which evolves over generations. Populations first respond with individual behavioural plasticity, whereafter changes may arise through innovations and the social transmission of behavioural patterns within and across generations, and, finally, by evolution of the behavioural response over generations. Only a restricted number of species show behavioural adaptations that make them thrive in severely disturbed environments. Hence, rapid human‐induced disturbances often decrease the diversity of native species, while facilitating the spread of invasive species with highly plastic behaviours. Consequently, behavioural responses to human‐induced environmental change can have profound effects on the distribution, adaptation, speciation and extinction of populations and, hence, on biodiversity. A better understanding of the mechanisms of behavioural responses and their causes and consequences could improve our ability to predict the effects of human‐induced environmental change on individual species and on biodiversity.     

Matching habitat choice promotes species persistence under climate change

Species may survive under contemporary climate change by either shifting their range or adapting locally to the warmer conditions. Theoretical and empirical studies recently underlined that dispersal, the central mechanism behind these responses, may depend on the match between an individuals’ phenotype and local environment. Such matching habitat choice is expected to induce an adaptive gene flow, but it now remains to be studied whether this local process could promote species’ responses to climate change. Here, we investigate this by developing an individual‐based model including either random dispersal or temperature‐dependent matching habitat choice. We monitored population composition and distribution through space and time under climate change. Relative to random dispersal, matching habitat choice induced an adaptive gene flow that lessened spatial range loss during climate warming by improving populations’ viability within the range (i.e. limiting range fragmentation) and by facilitating colonization of new habitats at the cold margin. The model even predicted range contraction under random dispersal but range expansion under optimal matching habitat choice. These benefits of matching habitat choice for population persistence mostly resulted from adaptive immigration decision and were greater for populations with larger dispersal distance and higher emigration probability. We also found that environmental stochasticity resulted in suboptimal matching habitat choice, decreasing the benefits of this dispersal mode under climate change. However population persistence was still better under suboptimal matching habitat choice than under random dispersal. Our results highlight the urgent need to implement more realistic mechanisms of dispersal such as matching habitat choice into models predicting the impacts of ongoing climate change on biodiversity.     

Multi-Axis Niche Examination of Ecological Specialization: Responses to Heat,Desiccation and Starvation Stress in Two Species of Pit-Building Antlions

Classical ecological studies discussing specialization usually focus on species’ performance along one niche axis. This approach may overlook niche differentiation evident in another dimension which could explain species co-occurrence. The present research exemplifies a comprehensive approach to examining local adaptation. Specifically, we examined multiple niche axes by subjecting a model organism to various experimental conditions to monitor responses to extreme stress associated with heat, desiccation and starvation. Our model system comprised two pit-building antlions: the habitat generalist Myrmeleon hyalinus and the habitat specialist Cueta lineosa. Previous research has shown that the foraging performance of the generalist is better than that of the specialist, even in the latter’s characteristic habitat. We illustrate that this apparent superiority of the habitat generalist does not manifest itself along other niche axes; rather, the habitat specialist holds a set of traits that provide an advantage under harsh environmental conditions. Specifically, C. lineosa has an advantage over M. hyalinus at high temperatures, exhibiting a higher survival rate and improved foraging success (i.e., high-temperature specialist). C. lineosa is also more efficient in its energy budget, losing less mass during starvation and gaining mass more efficiently during feeding. This superior efficiency is a result of physiological adaptations as well as behavioural responses to harsh conditions. In conclusion, our results imply that the habitat specialization of C. lineosa has not led it towards an evolutionary dead-end.     

Dynamics of a stochastic predator-prey model with habitat complexity and prey aggregation

Interplay between predator and prey is a complex process in ecosystems due to its nature. The population dynamics can be affected by many extrinsic and intrinsic factors. In this paper, we make an attempt to uncover the effects from environmental disturbances when populations are subject to habitat complexity and aggregation effect. We firstly propose a stochastic predator-prey model with habitat complexity and aggregation efficiency for prey. We then mathematically analyze the model, to demonstrate the existence, uniqueness and the stochastically ultimately boundedness of the global positive solution, and to establish sufficient conditions for the existence of ergodic stationary distribution of the solution. We also establish sufficient conditions under which either only predator population dies out or the entire predator-prey model becomes extinct. Our theoretical and numerical results indicate that: (1) the environmental noises are disadvantage for the survival of biological populations; (2) when the density of prey is greater than one, prey aggregation can heighten the capability of predator species to capture prey and reduce the effect of environmental fluctuations, while when the density of prey is less than one, the results are opposite; (3) habitat complexity is propitious to the survival of prey population and may seriously threaten the persistence of the predator population.     

Influence of climate change and postdelisting management on long‐term population viability of the conservation‐reliant Kirtland's Warbler

Rapid global climate change is resulting in novel abiotic and biotic conditions and interactions. Identifying management strategies that maximize probability of long‐term persistence requires an understanding of the vulnerability of species to environmental changes. We sought to quantify the vulnerability of Kirtland's Warbler (Setophaga kirtlandii), a rare Neotropical migratory songbird that breeds almost exclusively in the Lower Peninsula of Michigan and winters in the Bahamian Archipelago, to projected environmental changes on the breeding and wintering grounds. We developed a population‐level simulation model that incorporates the influence of annual environmental conditions on the breeding and wintering grounds, and parameterized the model using empirical relationships. We simulated independent and additive effects of reduced breeding grounds habitat quantity and quality, and wintering grounds habitat quality, on population viability. Our results indicated the Kirtland's Warbler population is stable under current environmental and management conditions. Reduced breeding grounds habitat quantity resulted in reductions of the stable population size, but did not cause extinction under the scenarios we examined. In contrast, projected large reductions in wintering grounds precipitation caused the population to decline, with risk of extinction magnified when breeding habitat quantity or quality also decreased. Our study indicates that probability of long‐term persistence for Kirtland's Warbler will depend on climate change impacts to wintering grounds habitat quality and contributes to the growing literature documenting the importance of considering the full annual cycle for understanding population dynamics of migratory species.     

Frequency distribution of coping strategies in four populations of brown trout (Salmo trutta)

In a challenging situation some animals respond by active avoidance, aggression and an activation of the sympathetic nervous system whereas others respond by immobility, low levels of aggression and a predominant adrenocortical stress response. When consistent over time and across situations such inter-individual differences in behavioural and physiological stress responses are referred to as stress coping strategies. In a previous study we reported the existence of two distinct stress coping strategies in a sea-ranched brown trout (Salmo trutta) population. Using the same method, we here show that four brown trout populations with different origin, but reared under identical conditions, differ in their endocrine stress response, behaviour during hypoxia and aggression. Further more, if individuals are classified as high- and low responsive based on post-stress blood plasma noradrenalin levels (indicator of sympathetic reactivity) the frequency distribution shows that populations with hatchery origin are biased towards having higher frequencies of high responsive individuals. However, the number of high responsive trout ranges from 14-48% in the different populations which shows that generally the frequency is biased towards lower levels of high responsive individuals. We discuss different frequency-dependent mechanisms that maintain multiple phenotypes in populations and speculate about differences in selection regime among the studied populations.     

Behavioural flexibility in the mating system buffers population extinction: lessons from the lesser spotted woodpecker Picoides minor

1. In most stochastic models addressing the persistence of small populations, environmental noise is included by imposing a synchronized effect of the environment on all individuals. However, buffer mechanisms are likely to exist that may counteract this synchronization to some degree. 2. We have studied whether the flexibility in the mating system, which has been observed in some bird species, is a potential mechanism counteracting the synchronization of environmental fluctuations. Our study organism is the lesser spotted woodpecker Picoides minor (Linnaeus), a generally monogamous species. However, facultative polyandry, where one female mates with two males with separate nests, was observed in years with male-biased sex ratio. 3. We constructed an individual-based model from data and observations of a population in Taunus, Germany. We tested the impact of three behavioural scenarios on population persistence: (1) strict monogamy; (2) polyandry without costs; and (3) polyandry assuming costs in terms of lower survival and reproductive success for secondary males. We assumed that polyandry occurs only in years with male-biased sex ratio and only for females with favourable breeding conditions. 4. Even low rates of polyandry had a strong positive effect on population persistence. The increase of persistence with carrying capacity was slower in the monogamous scenario, indicating strong environmental noise. In the polyandrous scenarios, the increase of persistence was stronger, indicating a buffer mechanism. In the polyandrous scenarios, populations had a higher mean population size, a lower variation in number of individuals, and recovered faster after a population breakdown. Presuming a realistic polyandry rate and costs for polyandry, there was still a strong effect of polyandry on persistence. 5. The results show that polyandry and in general flexibility in mating systems is a buffer mechanism that can significantly reduce the impact of environmental and demographic noise in small populations. Consequently, we suggest that even behaviour that seems to be exceptional should be considered explicitly when predicting the persistence of populations.     

Revisiting the Role of Individual Variability in Population Persistence and Stability

Populations often exhibit a pronounced degree of individual variability and this can be important when constructing ecological models. In this paper, we revisit the role of inter-individual variability in population persistence and stability under predation pressure. As a case study, we consider interactions between a structured population of zooplankton grazers and their predators. Unlike previous structured population models, which only consider variability of individuals according to the age or body size, we focus on physiological and behavioural structuring. We first experimentally demonstrate a high degree of variation of individual consumption rates in three dominant species of herbivorous copepods (Calanus finmarchicus, Calanus glacialis, Calanus euxinus) and show that this disparity implies a pronounced variation in the consumption capacities of individuals. Then we construct a parsimonious predator-prey model which takes into account the intra-population variability of prey individuals according to behavioural traits: effectively, each organism has a ‘personality’ of its own. Our modelling results show that structuring of prey according to their growth rate and vulnerability to predation can dampen predator-prey cycles and enhance persistence of a species, even if the resource stock for prey is unlimited. The main mechanism of efficient top-down regulation is shown to work by letting the prey population become dominated by less vulnerable individuals when predator densities are high, while the trait distribution recovers when the predator densities are low.     

Landscape genetics of an endangered lemur (Propithecus tattersalli) within its entire fragmented range

Habitat fragmentation may strongly reduce individuals’ dispersal among resource patches and hence influence population distribution and persistence. We studied the impact of landscape heterogeneity on the dispersal of the golden‐crowned sifaka (Propithecus tattersalli), an endangered social lemur species living in a restricted and highly fragmented landscape. We combined spatial analysis and population genetics methods to describe population units and identify the environmental factors which best predict the rates and patterns of genetic differentiation within and between populations. We used non‐invasive methods to genotype 230 individuals at 13 microsatellites in all the main forest fragments of its entire distribution area. Our analyses suggest that the Manankolana River and geographical distance are the primary structuring factors, while a national road crossing the region does not seem to impede gene flow. Altogether, our results are in agreement with a limited influence of forest habitat connectivity on gene flow patterns (except for North of the species’ range), suggesting that dispersal is still possible today among most forest patches for this species. Within forest patches, we find that dispersal is mainly among neighbouring social groups, hence confirming previous behavioural observations.     

Correlated extinctions, colonizations and population fluctuations in a highly connected ringlet butterfly metapopulation

 The persistence of metapopulations is likely to be highly dependent on whether population dynamics are correlated among habitat patches as a result of migration between patches and spatially-correlated environmental stochasticity (weather effects). We examined whether population dynamics of the ringlet butterfly, Aphantopus hyperantus, were synchronous in an area of approximately 0.5 km², with respect to extinction, colonization and population fluctuations. Monks Wood Butterfly Monitoring Scheme transect count data from 1973 to 1995, revealed (A) a major environmental perturbation, the drought of 1976, which caused synchronized extinctions of A. hyperantus in subsequent years, (B) synchronized recolonization in years following the large number of apparent extinctions, and (C) population changes by A. hyperantus were highly correlated in many of the 14 sections of the transect, presumably reflecting similar responses to environmental stochasticity, and the exchange of individuals among sections. However, extinction and population synchrony depended on habitat type. Following the 1976 drought, A. hyperantus apparently became extinct from the most open and most shady habitats it occupied, with some persistence in habitats of intermediate shading, thus showing retraction to core populations in central parts of an environmental gradient, albeit with an average shift to relatively open habitat. Populations at extreme ends of the environmental gradient occupied by A. hyperantus fluctuated least synchronously, suggesting a potential buffering effect of habitat heterogeneity, but this was not crucial to survival after the 1976 drought. Thus, not all habitats are equally important to persistence. Correlated temporal dynamics, variation in habitat quality and the interaction between habitat quality and temporal environmental stochasticity are important determinants of metapopulation persistence and should be incorporated in metapopulation models. Received: 26 April 1996 / Accepted: 17 July 1996     

Relictual physiological ecology in the threatened land snail Codringtonia helenae: A cause for decline in a changing environment?

Land snails often exhibit intra-annual cycles of activity interspersed by periods of dormancy (hibernation/aestivation), accompanied by a range of behavioural and physiological adaptations to ensure their survival under adverse environmental conditions. These adaptations are useful to understand species-specific habitat requirements and to predict their response to environmental changes. We examined the seasonal physiological and biochemical composition patterns of the threatened land snail Codringtonia helenae, endemic to Greece, in relation to its behavioural ecology and climatic conditions. Fuel reserves (glycogen, lipids, proteins) and water were accumulated prior to aestivation, but subsequently were rapidly depleted. LDH exhibited substantial rise during aestivation, suggesting that anaerobic pathways may provide additional energy. The major outcome of our study was the unambiguous discrimination of the four life-cycle periods. Most remarkable was the clear distinction of the aestivation period, with hibernation, the other dormancy period, showing similarity with the two active periods but not with aestivation. We observed disassociation between behavioural and physiological responses and climatic conditions. The physiological responses of C. helenae were effective during hibernation, but only partly compensate the effect of adverse conditions during aestivation, since its aestivating behaviour is occasional and time limited. Perhaps, the behavioural ecology of Codringtonia is relictual and shaped during past environmental conditions. This constitutes an important extinction threat considering the current climatic trends and the deterioration of the habitat of that species due to human activities.     

Climate mediates the success of migration strategies in a marine predator

Individual behavioural specialisation has far‐reaching effects on fitness and population persistence. Theory predicts that unconditional site fidelity, that is fidelity to a site independent of past outcome, provides a fitness advantage in unpredictable environments. However, the benefits of alternative site fidelity strategies driving intraspecific variation remain poorly understood and have not been evaluated in different environmental contexts. We show that contrary to expectation, strong and weak site fidelity strategies in migratory northern elephant seals performed similarly over 10 years, but the success of each strategy varied interannually and was strongly mediated by climate conditions. Strong fidelity facilitated stable energetic rewards and low risk, while weak fidelity facilitated high rewards and high risk. Weak fidelity outperformed strong fidelity in anomalous climate conditions, suggesting that the evolutionary benefits of site fidelity may be upended by increasing environmental variability. We highlight how individual behavioural specialisation may modulate the adaptive capacity of species to climate change.     

Integrodifference models for persistence in fragmented habitats

Integrodifference models of growth and dispersal are analyzed on finite domains to investigate the effects of emigration, local growth dynamics and habitat heterogeneity on population persistence. We derive the bifurcation structure for a range of population dynamics and present an approximation that allows straighforward calculation of the equilibrium populations in terms of local growth dynamics and dispersal success rates. We show how population persistence in a heterogeneous environment depends on the scale of the heterogeneity relative to the organism's characteristic dispersal distance. When organisms tend to disperse only a short distance, population persistence is dominated by local conditions in high quality patches, but when dispersal distance is relatively large, poor quality habitat exerts a greater influence.     

Habitat selection reduces extinction of populations subject to Allee effects

Theoretical studies indicate that a single population under an Allee effect will decline to extinction if reduced below a particular threshold, but the existence of multiple local populations connected by random dispersal improves persistence of the global population. An additional process that can facilitate persistence is the existence of habitat selection by dispersers. Using analytic and simulation models of population change, I found that when habitat patches exhibiting Allee effects are connected by dispersing individuals, habitat selection by these dispersers increases the likelihood that patches persist at high densities, relative to results expected by random settlement. Populations exhibiting habitat selection also attain equilibrium more quickly than randomly dispersing populations. These effects are particularly important when Allee effects are large and more than two patches exist. Integrating habitat selection into population dynamics may help address why some studies have failed to find extinction thresholds in populations, despite well-known Allee effects in many species.     

Effects of landscape and patch-level attributes on regional population persistence

Species responses are influenced by processes operating at multiple scales, yet many conservation studies and management actions are focused on a single scale. Although landscape-level habitat conditions (i.e., habitat amount, fragmentation and landscape quality) are likely to drive the regional persistence of spatially structured populations, patch-level factors (i.e., patch size, isolation, and quality) may also be important. To determine the spatial scales at which habitat factors influence the regional persistence of endangered Ord's kangaroo rats (Dipodomys ordii) in Alberta, Canada, we simulated population dynamics under a range of habitat conditions. Using a spatially-explicit population model, we removed groups of habitat patches based on their characteristics and measured the resulting time to extinction. We used proportional hazards models to rank the influence of landscape and interacting patch-level variables. Landscape quality was the most influential variable followed by patch quality, with both outweighing landscape- and patch-level measures of habitat quantity and fragmentation/proximity. Although habitat conservation and restoration priorities for this population should be in maximizing the overall quality of the landscape, population persistence depends on how this goal is achieved. Patch quality exerted a significant influence on regional persistence, with the removal of low quality road margin patches (sinks) reducing the risk of regional extinction. Strategies for maximizing overall landscape quality that omit patch-level considerations may produce suboptimal or detrimental results for regional population persistence, particularly where complex local population dynamics (e.g., source-sink dynamics) exist. This study contributes to a growing body literature that suggests that the prediction of species responses and future conservation actions may best be assessed with a multi-scale approach that considers habitat quality and that the success of conservation actions may depend on assessing the influences of habitat factors at multiple scales.     

Competition avoidance drives individual differences in response to a changing food resource in sticklebacks

Within the same population, individuals often differ in how they respond to changes in their environment. A recent series of models predicts that competition in a heterogeneous environment might promote between‐individual variation in behavioural plasticity. We tested groups of sticklebacks in patchy foraging environments that differed in the level of competition. We also tested the same individuals across two different social groups and while alone to determine the social environment's influence on behavioural plasticity. In support of model predictions, individuals consistently differed in behavioural plasticity when the presence of conspecifics influenced the potential payoffs of a foraging opportunity. Whether individuals maintained their level of behavioural plasticity when placed in a new social group depended on the environmental heterogeneity. By explicitly testing predictions of recent theoretical models, we provide evidence for the types of ecological conditions under which we would expect, and not expect, variation in behavioural plasticity to be favoured.     

Fluctuating asymmetry and herbivory in two ontogenetical stages of Chamaecrista semaphora in restored and natural environments

Reintroduced environments represent stressful conditions to plants that can be observed in different ways. We evaluated the relationships between fluctuating asymmetry (FA), herbivory, and plant ontogeny of Chamaecrista semaphora (Fabaceae) under natural and restored habitat conditions. The patterns of leaf FA and herbivory by folivorous insects (chewing) on saplings and mature plants in each habitat were determined. No relationship was found between FA and herbivory on the two ontogenetic stages in both environments, suggesting that FA did not represent an indicator of stress. The frequency and amount of leaf area removed by folivores were higher in saplings compared to adult plants under the natural habitat, while the opposite trend was observed on restored habitat for adult plants. The restored habitat did not represent an environmental stress condition to C. semaphora, indicating that this endemic plant may represent a good candidate to restoration programs in harsh environments. However, we observed great differences in leaf FA and herbivory among individuals within habitats. Knowledge on plant quality, competition, physiology, and interactions with natural enemies are highly needed to support long lasting programmes on restoration of harsh environments.     

Individual‐based modelling of resource competition to predict density‐dependent population dynamics: a case study with white storks

Density regulation influences population dynamics through its effects on demographic rates and consequently constitutes a key mechanism explaining the response of organisms to environmental changes. Yet, it is difficult to establish the exact form of density dependence from empirical data. Here, we developed an individual‐based model to explore how resource limitation and behavioural processes determine the spatial structure of white stork Ciconia ciconia populations and regulate reproductive rates. We found that the form of density dependence differed considerably between landscapes with the same overall resource availability and between home range selection strategies, highlighting the importance of fine‐scale resource distribution in interaction with behaviour. In accordance with theories of density dependence, breeding output generally decreased with density but this effect was highly variable and strongly affected by optimal foraging strategy, resource detection probability and colonial behaviour. Moreover, our results uncovered an overlooked consequence of density dependence by showing that high early nestling mortality in storks, assumed to be the outcome of harsh weather, may actually result from density dependent effects on food provision. Our findings emphasize that accounting for interactive effects of individual behaviour and local environmental factors is crucial for understanding density‐dependent processes within spatially structured populations. Enhanced understanding of the ways animal populations are regulated in general, and how habitat conditions and behaviour may dictate spatial population structure and demographic rates is critically needed for predicting the dynamics of populations, communities and ecosystems under changing environmental conditions.     

Inertia: the discrepancy between individual and common good in dispersal and prospecting behaviour

The group selection debate of the 1960s made it clear that evolution does not necessarily increase population performance. Individuals can be selected to have traits that diminish a common good and make population persistence difficult. At the extreme, the discrepancy between levels of selection is predicted to make traits evolve towards values at which a population can no longer persist (evolutionary suicide). Dispersal and prospecting are prime examples of traits that have a strong influence on population persistence under environmental and demographic stochasticity. Theory predicts that an ‘optimal’ dispersal strategy from a population point of view can differ considerably from that produced by individual‐level selection. Because dispersal is frequently risky or otherwise costly, individuals are often predicted to disperse less than would be ideal for population performance (persistence or size). We define this discrepancy as ‘inertia’ and examine current knowledge of its occurrence and effects on population dynamics in nature. We argue that inertia is potentially widespread but that a framework is currently lacking for predicting precisely the extent to which it has a real influence on population persistence. The opposite of inertia, ‘hypermobility’ (more dispersal by individuals than would maximize population performance) remains a possibility: it is known that highest dispersal rates do not lead to best expected population performance, and examples of such high dispersal evolving exist at least in the theoretical literature. We also show, by considering prospecting behaviour, that similar issues arise in species with advanced cognitive and learning abilities. Individual prospecting strategies and the information acquired during dispersal are known to influence the decisions and therefore the fate of individuals and, as a corollary, populations. Again, the willingness of individuals to sample environments might evolve to levels that are not optimal for populations. This conflict can take intriguing forms. For example, better cognitive abilities of individuals may not always lead to better population‐level performance. Simulation studies have found that ‘blind’ dispersal can lead to better connected metapopulations than cognitively more advanced habitat choice rules: the latter can lead to too many individuals sticking to nearby safe habitat. The study of the mismatch between individual and population fitness should not be a mere intellectual exercise. Population managers typically need to take a population‐level view of performance, which may necessitate human intervention if it differs from what is selected for. We conclude that our knowledge of inertia and hypermobility would advance faster if theoretical studies—without much additional effort—quantified the population consequences of the evolving traits and compared this with hypothetical (not selectively favoured) dispersal rules, and if empirical studies were similarly conducted with the differing levels of selection in mind.     

Linking trade-offs in habitat selection with the occurrence of functional responses for moose living in two nearby study areas

A species may modify its relative habitat use with changing availability, generating functional responses in habitat selection. Functional responses in habitat selection are expected to occur when animals experience trade-offs influencing their habitat selection, but only a few studies to date have explicitly linked functional responses to the underlying trade-offs faced by the animals. We used data from 39 female moose fitted with GPS telemetry collars in two nearby study areas in Canada to investigate if moose (1) were faced with a food/cover trade-off in habitat selection, as typically acknowledged in the literature, and (2) showed a functional response in their use of food/cover-rich habitats. We also examined how habitat selection patterns varied seasonally, and between study areas. The occurrence of functional responses varied strongly between study areas, and could not always be related to a measurable food/cover trade-off. Functional responses were observed more often in the study area where the environmental conditions were more severe (colder temperatures, higher precipitations, and lower food availability). Selection coefficients were also less variable among individuals in that study area, suggesting that severe environmental conditions may constrain individuals to a few selection tactics and promote the development of functional responses. Moose reacted to the availability of different habitat types in different seasons, reflecting the changing trade-offs faced by the animals. We found considerable behavioral differences between individuals from two adjacent study areas, and therefore recommend caution when extrapolating habitat selection results. We advocate for the wider use of functional responses to identify critical habitats for a species from a management or conservation perspective.