Habitat loss and fragmentation affecting mammal and bird communities—The role of interspecific competition and individual space use

Fragmentation and loss of habitat are major threats to animal communities and are therefore important to conservation. Due to the complexity of the interplay of spatial effects and community processes, our mechanistic understanding of how communities respond to such landscape changes is still poor. Modelling studies have mostly focused on elucidating the principles of community response to fragmentation and habitat loss at relatively large spatial and temporal scales relevant to metacommunity dynamics. Yet, it has been shown that also small scale processes, like foraging behaviour, space use by individuals and local resource competition are also important factors. However, most studies that consider these smaller scales are designed for single species and are characterized by high model complexity. Hence, they are not easily applicable to ecological communities of interacting individuals. To fill this gap, we apply an allometric model of individual home range formation to investigate the effects of habitat loss and fragmentation on mammal and bird communities, and, in this context, to investigate the role of interspecific competition and individual space use. Results show a similar response of both taxa to habitat loss. Community composition is shifted towards higher frequency of relatively small animals. The exponent and the 95%-quantile of the individual size distribution (ISD, described as a power law distribution) of the emerging communities show threshold behaviour with decreasing habitat area. Fragmentation per se has a similar and strong effect on mammals, but not on birds. The ISDs of bird communities were insensitive to fragmentation at the small scales considered here. These patterns can be explained by competitive release taking place in interacting animal communities, with the exception of bird's buffering response to fragmentation, presumably by adjusting the size of their home ranges. These results reflect consequences of higher mobility of birds compared to mammals of the same size and the importance of considering competitive interaction, particularly for mammal communities, in response to landscape fragmentation. Our allometric approach enables scaling up from individual physiology and foraging behaviour to terrestrial communities, and disentangling the role of individual space use and interspecific competition in controlling the response of mammal and bird communities to landscape changes.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.     

Linking foraging decisions to residential yard bird composition

Urban bird communities have higher densities but lower diversity compared with wildlands. However, recent studies show that residential urban yards with native plantings have higher native bird diversity compared with yards with exotic vegetation. Here we tested whether landscape designs also affect bird foraging behavior. We estimated foraging decisions by measuring the giving-up densities (GUD; amount of food resources remaining when the final forager quits foraging on an artificial food patch, i.e seed trays) in residential yards in Phoenix, AZ, USA. We assessed how two yard designs (mesic: lush, exotic vegetation; xeric: drought-tolerant and native vegetation) differed in foraging costs. Further, we developed a statistical model to calculate GUDs for every species visiting the seed tray. Birds foraging in mesic yards depleted seed trays to a lower level (i.e. had lower GUDs) compared to birds foraging in xeric yards. After accounting for bird densities, the lower GUDs in mesic yards appeared largely driven by invasive and synanthropic species. Furthermore, behavioral responses of individual species were affected by yard design. Species visiting trays in both yard designs had lower GUDs in mesic yards. Differences in resource abundance (i.e., alternative resources more abundant and of higher quality in xeric yards) contributed to our results, while predation costs associated with foraging did not. By enhancing the GUD, a common method for assessing the costs associated with foraging, our statistical model provided insights into how individual species and bird densities influenced the GUD. These differences we found in foraging behavior were indicative of differences in habitat quality, and thus our study lends additional support for native landscapes to help reverse the loss of urban bird diversity.     

Fast food in ant communities: how competing species find resources

An understanding of foraging behavior is crucial to understanding higher level community dynamics; in particular, there is a lack of information about how different species discover food resources. We examined the effect of forager number and forager discovery capacity on food discovery in two disparate temperate ant communities, located in Texas and Arizona. We defined forager discovery capacity as the per capita rate of resource discovery, or how quickly individual ants arrived at resources. In general, resources were discovered more quickly when more foragers were present; this was true both within communities, where species identity was ignored, as well as within species. This pattern suggests that resource discovery is a matter of random processes, with ants essentially bumping into resources at a rate mediated by their abundance. In contrast, species that were better discoverers, as defined by the proportion of resources discovered first, did not have higher numbers of mean foragers. Instead, both mean forager number and mean forager discovery capacity determined discovery success. The Texas species used both forager number and capacity, whereas the Arizona species used only forager capacity. There was a negative correlation between a species’ prevalence in the environment and the discovery capacity of its foragers, suggesting that a given species cannot exploit both high numbers and high discovery capacity as a strategy. These results highlight that while forager number is crucial to determining time to discovery at the community level and within species, individual forager characteristics influence the outcome of exploitative competition in ant communities.     

The consequences of size dependent foraging for food web topology

A general question in biology is how processes at one scale, for example that of individual organisms, influence patterns at larger scales, for example communities of interacting individuals. Here we ask how changing the size‐dependence of the foraging behaviour of individuals can influence the structure of food webs. We assembled communities using a model in which species interactions are determined by allometric foraging rules of (1) handling time and (2) attack rates, and also (3) the distribution of body sizes. We systematically varied these three factors and examined their effects on three community level, food web allometries: the generality ‐ mass correlation, the vulnerability ‐ mass correlation and the trophic height ‐ mass correlation. The results demonstrate how allometries of individual foraging behaviour (handling time and attack rates) are linked across scales of organisation: different community level allometries are influenced by different individual level allometries. For example, generality allometries in the community are most affected by the individual allometric relationships of the attack rate, whereas trophic level allometries in the community are more strongly influenced by variation in individual handling time allometries. Importantly, we also find that the shape of the body size distribution from which species are drawn has a substantial influence on how these links between scales operate. This study suggests that understanding the variation of size structure among ecological networks requires knowledge about the causes of variation in individual foraging behaviour and determinants of the regional body size distribution.     

At-sea distribution and scale-dependent foraging behaviour of petrels and albatrosses: a comparative study

1. In order to study and predict population distribution, it is crucial to identify and understand factors affecting individual movement decisions at different scales. Movements of foraging animals should be adjusted to the hierarchical spatial distribution of resources in the environment and this scale-dependent response to environmental heterogeneity should differ according to the forager's characteristics and exploited habitats. 2. Using First-Passage Time analysis, we studied scales of search effort and habitat used by individuals of seven sympatric Indian Ocean Procellariiform species fitted with satellite transmitters. We characterized their search effort distribution and examined whether species differ in scale-dependent adjustments of their movements according to the marine environment exploited. 3. All species and almost all individuals (91% of 122 individuals) exhibited an Area-Restricted Search (ARS) during foraging. At a regional scale (1000s km), foraging ranges showed a large spatial overlap between species. At a smaller scale (100s km, at which an increase in search effort occurred), a segregation in environmental characteristics of ARS zones (where search effort is high) was found between species. 4. Spatial scales at which individuals increased their search effort differed between species and also between exploited habitats, indicating a similar movement adjustment for predators foraging in the same habitat. ARS zones of the two populations of wandering albatross Diomedea exulans (Crozet and Kerguelen) were similar in their adjustments (i.e. same ARS scale) as well as in their environmental characteristics. These two populations showed a weak spatial overlap in their foraging distribution, with males foraging in more southerly waters than females in both populations. 5. This study demonstrates that predators of several species adjust their foraging behaviour to the heterogeneous environment and these scale-dependent movement adjustments depend on both forager and environment characteristics.     

State‐space modeling reveals habitat perception of a small terrestrial mammal in a fragmented landscape

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Is foraging in the desert ant,Messor pergandei (Hymenoptera: Formiceidae), limited by water?

Abstract.

• 1 The seed-harvesting ant Messor (Veromessor) prrgandei (Mayr) is a common inhabitant of southwestern deserts of the U.S.A. Foragers vary in size from less than 1 mg to more than 10 mg in body mass and may travel over 80 m on a single foraging trip. Their small size, long foraging range, and hot, arid habitat suggest that water stress may limit foraging activity. We examined intercolony and interindividual variation in water loss of M.pergandei foragers under several different situations in the field.
• 2 Colonies differed significantly in minimum critical water content (W_c) of individual foragers (water content below which foragers are incapable of normal locomotion). In one colony small workers had disproportionately higher W_c than larger workers; in the other colony W_c was isometric with body size.
• 3 Groups of workers confined in the field approached W_c only after normal foraging stopped and substrate temperatures exceeded 45°C, while water content of individual foragers did not approach the W_c during normal foraging periods. Moreover, seed load and distance travelled did not negatively affect forager water content, as measured on return to the nest: indeed, our results suggest that forager hydration level may influence load selection and/or foraging distance. We conclude that, under normal circumstances, foraging in M.pergandei is not water-limited.

     

Patch use, apprehension, and vigilance behavior of Nubian Ibex under perceived risk of predation

Foraging theory predicts that animals will sacrifice feedingeffort in order to reduce predation risk. Once a forager choosesa habitat, it must decide how to allocate its foraging effort.Nubian Ibex are diurnal, social, cliff-dwelling herbivores.Many of their characteristics seem to have evolved as responsesto predation risk. In order to assess the effects that perceivedrisk of predation might have on foraging behavior of free-rangingNubian Ibex in the Negev Desert, Israel, we measured giving-updensities (GUDs) in artificial food patches and used them togauge apprehension level. (Apprehension can be defined as areduction in attention devoted to performing an activity asa consequence of reallocating attention to detecting or respondingto predation risk. A forager can also be vigilant. Vigilanceis often defined as time spent scanning the surroundings withthe head up.) We also quantified time budgeting using focalobservation of individual Nubian Ibex. Habitat preferences andpatch selectivity as a measure of apprehension were considered.In particular, we tested the effect of distance from refugeon GUDs, the effect of micropatch structure on selectivity,and the effect of distance from the refuge and group size onNubian Ibex vigilance level and apprehension. Nubian Ibex allocatetheir foraging effort more toward patches closer to the escapeterrain. At the same time, Nubian Ibex are more apprehensiveat intermediate distances from the cliff edge than nearer thecliff, and their use of vigilance increases with distance fromthe cliff edge. These results suggest that Nubian Ibex may switchfrom apprehension to a more extreme behavior of vigilance atgreater distances from the refuge. This study demonstrated theuse of antipredatory behaviors, apprehension, and vigilanceby a forager. Estimating apprehension and vigilance levels ofa forager simultaneously gives a more complete and accuratepicture of how the habitat is perceived by them and combinedwith measurements of GUD allow a more accurate assessment ofhabitat quality.     

Modeling the emergence of migratory corridors and foraging hot spots of the green sea turtle

Environmental factors shape the spatial distribution and dynamics of populations. Understanding how these factors interact with movement behavior is critical for efficient conservation, in particular for migratory species. Adult female green sea turtles, Chelonia mydas, migrate between foraging and nesting sites that are generally separated by thousands of kilometers. As an emblematic endangered species, green turtles have been intensively studied, with a focus on nesting, migration, and foraging. Nevertheless, few attempts integrated these behaviors and their trade‐offs by considering the spatial configurations of foraging and nesting grounds as well as environmental heterogeneity like oceanic currents and food distribution. We developed an individual‐based model to investigate the impact of local environmental conditions on emerging migratory corridors and reproductive output and to thereby identify conservation priority sites. The model integrates movement, nesting, and foraging behavior. Despite being largely conceptual, the model captured realistic movement patterns which confirm field studies. The spatial distribution of migratory corridors and foraging hot spots was mostly constrained by features of the regional landscape, such as nesting site locations, distribution of feeding patches, and oceanic currents. These constraints also explained the mixing patterns in regional forager communities. By implementing alternative decision strategies of the turtles, we found that foraging site fidelity and nesting investment, two characteristics of green turtles' biology, are favorable strategies under unpredictable environmental conditions affecting their habitats. Based on our results, we propose specific guidelines for the regional conservation of green turtles as well as future research suggestions advancing spatial ecology of sea turtles. Being implemented in an easy to learn open‐source software, our model can coevolve with the collection and analysis of new data on energy budget and movement into a generic tool for sea turtle research and conservation. Our modeling approach could also be useful for supporting the conservation of other migratory marine animals.     

Behavioural responses of the lizard Pedioplanis l. lineoocellata to overgrazing

We studied the effects of overgrazing on the foraging behaviour of the lizard Pedioplanis l. lineoocellata (Spotted Sand Lizard), a sit-and-wait forager, in habitats of differing vegetation states to determine the effects of habitat degradation on this species. At high grazing intensity where vegetation cover and diversity is low, the lizard P. lineoocellata moves more frequently, spends more time moving and covers larger distances than in habitats where vegetation cover and diversity is high. These behavioural changes in movement patterns can be explained by less abundant prey in habitats with low vegetation cover and diversity. Although morphology, phylogeny and physiology of P. lineoocellata should constrain the change in foraging behaviour, the species has modified its foraging strategy from sit-and-wait to actively foraging. We assume that this behavioural flexibility of P. lineoocellata is a buffer mechanism enabling the species to use and survive in degraded (unfavourable) habitats.     

Population and community consequences of spatial subsidies derived from central-place foraging

Central-place foragers, such as ants, beavers, and colonial seabirds, can act as biological conduits, subsidizing local communities with allochthonous resources. To explore the consequences of such biologically vectored resource redistribution, we draw on an example from cave ecology and develop a population-level model of central-place foraging based on the dispersal kernel framework. We explore how the size of the patch in which central-place foraging occurs and the spatial distribution of foragers within that patch feed back to influence the population dynamics of the central-place forager and the species richness of the associated recipient community. We demonstrate that the particular way in which a population of central-place foragers uses space has two important effects. First, space use determines the stability of the forager population and establishes patch size thresholds for persistence, stable equilibria, and limit cycles. Second, alternative foraging kernels lead to qualitatively different scaling relationships between the size of the foraging patch and species richness back at the central place. These analyses provide a new link among elements of ecology related to animal behavior, population dynamics, and species diversity while also providing a novel perspective on the utility of integrodifference equations for problems in spatial ecology.     

Spatial and trophic niche differentiation in two sympatric lizards (Tropidurus torquatus and Cnemidophorus ocellifer) with different foraging tactics

Abstract In a ‘restinga’ habitat of southeastern Brazil, we studied the food habits and the microhabitat use of two lizards with distinct foraging modes: the tropidurid Tropidurus torquatus, a sit-and-wait predator, and the teiid Cnemidophorus ocellifer, a wide forager. The diet of the two species differed strongly, indicating a low level of similarity in their trophic niche. The sit-and-wait predator fed mainly on mobile prey, whereas the wide forager fed mainly on sedentary prey (larvae). The spatial niche breadth of T. torquatus was larger than that of C. ocellifer. Despite interspecific differences, the two species overlapped greatly in micro-habitat use. The data indicate that at Linhares the two lizard species differed more in food resources than in microhabitat, and that most of the food differences reflect the foraging patterns of the species.     

Environmental heterogeneity amplifies behavioural response to a temporal cycle

Resource acquisition is integral to maximise fitness, however in many ecosystems this requires adaptation to resource abundance and distributions that seldom stay constant. For predators, prey availability can vary at fine spatial and temporal scales as a result of changes in the physical environment, and therefore selection should favour individuals that can adapt their foraging behaviour accordingly. The tidal cycle is a short, yet predictable, temporal cycle, which can influence prey availability at temporal scales relevant to movement decisions. Here, we ask whether black‐legged kittiwakes Rissa tridactyla can adjust their foraging habitat selection according to the tidal cycle using GPS tracking studies at three sites of differing environmental heterogeneity. We used a hidden Markov model to classify kittiwake behaviour, and analysed habitat selection during foraging. As expected for a central‐place forager, we found that kittiwakes preferred to forage nearer to the breeding colony. However, we also show that habitat selection changed over the 12.4‐h tidal cycle, most likely because of changes in resource availability. Furthermore, we observed that environmental heterogeneity was associated with amplified changes in kittiwake habitat selection over the tidal cycle, potentially because environmental heterogeneity drives greater resource variation. Both predictable cycles and environmental heterogeneity are ubiquitous. Our results therefore suggest that, together, predictable cycles and environmental heterogeneity may shape predator behaviour across ecosystems.     

Multimodality in the individual size distributions of bird communities

Aim Body size often plays a significant role in community assembly through its impacts on the life history and ecological attributes of species. Insight into the importance of size in structuring communities can be gained by examining the distribution of sizes of individuals [i.e. the individual size distribution (ISD) or size spectrum] in a community. ISDs have been studied extensively in aquatic and tree communities, but have received little attention in terrestrial animal communities. Here, we conduct the first macroecological analysis of ISDs in terrestrial animal communities to determine whether they show broad‐scale consistency in shape. Location North America, north of Mexico. Methods Using likelihood‐based methods and Gaussian mixture modelling, coupled with data from the Breeding Bird Survey and Christmas Bird Count, we determine whether the ISDs for thousands of breeding and wintering North American bird communities are: (1) monotonically decreasing, (2) unimodal or (3) multimodal. Results We find that avian ISDs are consistently multimodal, with most characterized by more than five modes in both breeding and wintering communities from local to continental scales. In addition, the positions of these modes along the size axis are remarkably consistent. Main conclusions The striking consistency in the ISD within bird communities, as with tree and aquatic communities, indicates that the ISD is an important and informative characterization of resource utilization within an ecological assemblage. The differences in shape of the ISD among these groups also suggest that differences in body size‐related constraints affect interactions within a group and with the environment. Our results confirm that avian assemblages do exhibit structure along the body size axis, and therefore it will be fruitful to explore this pattern in greater detail.     

Foraging by forest ants under experimental climatic warming: a test at two sites

Climatic warming is altering the behavior of individuals and the composition of communities. However, recent studies have shown that the impact of warming on ectotherms varies geographically: species at warmer sites where environmental temperatures are closer to their upper critical thermal limits are more likely to be negatively impacted by warming than are species inhabiting relatively cooler sites. We used a large‐scale experimental temperature manipulation to warm intact forest ant assemblages in the field and examine the impacts of chronic warming on foraging at a southern (North Carolina) and northern (Massachusetts) site in eastern North America. We examined the influence of temperature on the abundance and recruitment of foragers as well as the number of different species observed foraging. Finally, we examined the relationship between the mean temperature at which a species was found foraging and the critical thermal maximum temperature of that species, relating functional traits to behavior. We found that forager abundance and richness were related to the experimental increase in temperature at the southern site, but not the northern site. Additionally, individual species responded differently to temperature: some species foraged more under warmer conditions, whereas others foraged less. Importantly, these species‐specific responses were related to functional traits of species (at least at the Duke Forest site). Species with higher critical thermal maxima had greater forager densities at higher temperatures than did species with lower critical thermal maxima. Our results indicate that while climatic warming may alter patterns of foraging activity in predictable ways, these shifts vary among species and between sites. More southerly sites and species with lower critical thermal maxima are likely to be at greater risk to ongoing climatic warming.     

Effect of local spatial plant distribution and conspecific density on bumble bee foraging behaviour

1. Size variations in pollinator populations may modify competitive interactions among foragers. Competition among pollinators has been shown to lead to one of two contrasting behaviours: either specialisation to the most profitable plant species or generalisation to several species. When foraging, pollinators are also confronted with heterogeneity in the spatial distribution of plant resources. Because variations in both the forager density and plant spatial distribution can affect pollinator behaviour, focus was on the interactive effect of these two factors. 2. Bumble bee (Bombus terrestris L.) individuals were trained on a focal species (Lotus corniculatus L.) and experimentally tested whether variations in the forager density (two or six bumble bees foraging together), plant community spatial distribution (two plant species: L. corniculatus and Medicago sativa, which were either patchily or randomly distributed), and their interaction modified bumble bee foraging behaviour. 3. It was shown that when confronted with a high forager density, bumble bees focused their visits towards the most familiar species, especially when foraging under a random plant distribution. These modifications affected the fruiting of the focal plant species, with a significantly lower reproductive success under low density/patchy conditions. 4. This study demonstrates that the foraging decisions of bumble bees are influenced by variations in both the conspecific density and plant spatial distribution. Given the increasing impact of human activities on plant‐pollinator communities, this raises the question of the potential implications of these results for plant communities in natural conditions when confronted with variations in the pollinator density and spatial distribution of plants.     

Landscape geometry determines community response to disturbance

Ecological communities are impacted by anthropogenic changes in both habitat geometry (i.e. amount, shape, fragmentation and connectivity of habitat) and disturbance regime. Although the effect of each of these drivers on diversity is well-documented, few studies have considered how habitat geometry and disturbance interact to affect diversity. We used a miniature landscape of moss patches to experimentally manipulate both habitat geometry and disturbance frequency on microarthropod communities. Species richness and abundance in local patches declined linearly with disturbance rate in all experimental landscapes, but the speed of this decline (a measure of ecological resilience) depended on the size and connectivity of the surrounding region. Reductions in region size had little effect on community resilience to disturbance until habitat loss resulted in complete loss of connectivity between patches, suggesting a threshold in community response to habitat loss. Beyond this threshold, repeated disturbance resulted in rapid declines in patch species richness and abundance and substantial changes in community composition. These effects of habitat geometry and disturbance on diversity were scale-dependent. Gamma (regional-scale) diversity was unaffected by habitat geometry, suggesting experimental reductions in alpha (local-scale) diversity were offset by increases in beta diversity. There was no effect of body size, abundance, or trophic position in determining species response to disturbance. Taxonomic grouping had a weak effect, with oribatids less affected by drought. We conclude that, in this system, dispersal from the surrounding metacommunity is vital in allowing recovery of local communities from disturbance. When habitat loss and fragmentation disrupt this process, extinctions result. Studies that examine separately the effects of habitat alterations and disturbance on diversity may therefore underestimate their combined effects.     

The Trail Less Traveled: Individual Decision-Making and Its Effect on Group Behavior

Social insect colonies are complex systems in which the interactions of many individuals lead to colony-level collective behaviors such as foraging. However, the emergent properties of collective behaviors may not necessarily be adaptive. Here, we examine symmetry breaking, an emergent pattern exhibited by some social insects that can lead colonies to focus their foraging effort on only one of several available food patches. Symmetry breaking has been reported to occur in several ant species. However, it is not clear whether it arises as an unavoidable epiphenomenon of pheromone recruitment, or whether it is an adaptive behavior that can be controlled through modification of the individual behavior of workers. In this paper, we used a simulation model to test how symmetry breaking is affected by the degree of non-linearity of recruitment, the specific mechanism used by individuals to choose between patches, patch size, and forager number. The model shows that foraging intensity on different trails becomes increasingly asymmetric as the recruitment response of individuals varies from linear to highly non-linear, supporting the predictions of previous work. Surprisingly, we also found that the direction of the relationship between forager number (i.e., colony size) and asymmetry varied depending on the specific details of the decision rule used by individuals. Limiting the size of the resource produced a damping effect on asymmetry, but only at high forager numbers. Variation in the rule used by individual ants to choose trails is a likely mechanism that could cause variation among the foraging behaviors of species, and is a behavior upon which selection could act.     

Effects of fragmented breeding habitat and resource distribution on behavior and survival of the bank vole (Myodes glareolus)

Habitat fragmentation causes negative population trends or even local extinction in many species. Understanding the role of fragmentation on behavior and space use of animals is an essential part of revealing the mechanism behind observed population declines. We studied experimentally the effects of small-scale habitat fragmentation on the distribution and movement of bank voles (Myodes glareolus) in seminatural enclosures. We predicted that besides habitat structure, availability of two resources, food for both sexes, and receptive females for males, determine individual distribution. We manipulated female density (either 0, 4, or 12 per enclosure) and followed movements of radio-collared males. We also studied the effect of unequal food distribution between the fragments on vole distribution. The habitat of 0.25-ha enclosures was manipulated by forming either one large, two medium-sized, or four small habitat patches surrounded by an inhospitable matrix. Female density and distribution clearly affected male spacing behavior. Males had larger home ranges in the medium female density enclosures. Furthermore, the use of the inhospitable and risky matrix area increased with habitat fragmentation. Food supplementation improved individual condition measured as body mass but did not affect breeding success. Our experiments demonstrated that both small-scale habitat fragmentation and resource distribution affect the behavior and condition of individuals. Increased fragmentation led to increased risk taking in both mating and foraging behavior. This should have direct survival and fitness consequences, and therefore our results may be extrapolated to population-level consequences of habitat fragmentation.