Carbon and nitrogen isotopes trace nutrient exchange in an ant-plant mutualism

By combining and reanalysing data from two independent field experiments we explore whether food limitation at the brood stage affects habitat selection in nesting mallards (Anas platyrhynchos). In an introduction experiment we found that, independent of treatment, some study lakes remained empty of wild mallard pairs (”empty lakes”), whereas on other lakes introduced birds attracted wild mallards (”attractive lakes”). In the other experiment we used mallard ducklings to address brood-stage food limitation by studying mass change of ducklings. We found that ducklings foraging on lakes that did not attract wild mallard pairs in the introduction experiment gained much less mass than those foraging on attractive lakes. In most cases ducklings even lost mass in the empty-lake foraging trials, providing strong evidence for food limitation. Therefore, lakes that remained empty of wild mallard pairs in the introduction experiment proved to be inferior brood habitats, particularly in terms of food. Our results give insight into the mechanisms underlying the general habitat selection hypotheses, specifically the ideal preemptive and conspecific attraction rules. The results further support our earlier conclusion that mallards do not use the ideal preemptive rule when selecting nesting lakes. However, conspecific attraction may not be generally applicable either, because, independent of the presence of introduced conspecifics, wild mallards somehow anticipated the low quality of the empty lakes as brood-rearing habitats and made their habitat-selection decision accordingly. Received: 22 March 1999 / Accepted: 7 October 1999     

Dynamics of habitat distribution in breeding mallards: assessing the applicability of current habitat selection models

The applicability of ideal free, ideal despotic and ideal preemptive habitat selection models to explain dynamics of habitat distribution of breeding mallards ( Anas platyrhynchos ) was explored. Data from 35 lakes studied between 1985 and 2000 were used to examine overall habitat distribution of breeding pairs, breeding success in different habitats, within-year order of habitat occupation, and density dependence of habitat distribution and breeding success. Two habitat types, rich and poor, were defined based on the structure and luxuriance of shore vegetation; each lake belonged to one or the other of the habitat types. Breeding pairs used the rich habitat more than expected, breeding density also being higher there than in the poor habitat. Both average brood density and breeding success were higher in the rich habitat than in the poor. Breeding success was not density dependent, neither when analysed separately for the habitat types, nor in the study area in general. Within season, arriving mallard pairs did not occupy rich lakes earlier than poor lakes. An isodar analysis based on between-year variation of the breeding density in rich and poor habitats revealed that habitat distribution of breeding pairs was not density dependent. By contrast, density in the rich habitat increased and proportional use of the poor habitat decreased with increasing overall population density, i.e. the rich habitat got increasingly crowded. None of the habitat selection models considered was applicable to explain the dynamics of habitat distribution of breeding mallards.     

Aggregations from using inadvertent social information: a form of ideal habitat selection

Social information in breeding site selection has received extensive study; however, few attempts have been made to link this process to pre‐existing models. We examine the importance of social information to three pertinent models of habitat selection that describe breeding aggregations and spatial patterns: 1) the ideal despotic distribution (IDD) which considers conspecific competition and habitat availability, 2) the perceptual constraints model which accounts for patch selection when animals experience a threshold of undetectable difference in quality, and 3) the “neighbourhood model” which predicts concordance between resources and settlers can be disrupted by conspecific attraction when resources are patchy. These models all predict initial settlers will select a high quality patch first. However, their predictions of subsequent settlement behaviour in remaining patches differ: the IDD predicts subsequent settlers will be distributed regularly, the perceptual constraints model predicts a random distribution, and the neighbourhood model predicts clustering from conspecific attraction. We examined which model best described settlement patterns of bobolink Dolichonyx oryzivorus and savannah sparrow Passerculus sandwichensis, in the context of social information. We observed settlement timing, quantified available resources, and determined where they occurred in the highest (local population “core”) and lowest densities (local population “periphery”). We then assessed whether individuals in the periphery settled in greater concordance with resources or conspecific presence. Core territories were clustered strongly on relevant resources, and these territory holders were older than in the periphery. Peripheral territories were likewise clustered but did not always co‐occur with the best available resources, matching the neighbourhood model prediction that social information may not always direct them to the best sites available. This suggests older individuals used their own experience to locate ideal habitat, whereas younger individuals attempted to aggregate on seemingly ideal habitat by using conspecific location; such information asymmetry due to age can be viewed as an “ideal aggregative distribution”.     

Wood warblers copy settlement decisions of poor quality conspecifics: support for the tradeoff between the benefit of social information use and competition avoidance

Social information use in songbird habitat selection commonly involves a conspecific attraction strategy. Individuals copy the breeding‐site choices of conspecifics, that is, bias their own settlement decisions towards sites (tracts of spatially limited habitat with similar structure) already occupied by others. In order to be adaptive, social information use has to be discriminative. Especially the decisions of good quality individuals, i.e. measuring high at observable fitness correlates, should be copied more frequently than those of poor quality individuals. It is unknown, however, whether songbirds discriminatively use conspecific presence by evaluating the quality of information providers in habitat selection. We experimentally tested whether wood warblers Phylloscopus sibilatrix selectively copied settlement decisions of conspecifics in relation to the quality of observed individuals. We also tested whether the use of social cues was influenced by the population density at a particular site in the preceding year. We found that wood warblers selectively used intraspecific social information, but in a pattern opposite to that expected based on existing hypotheses. Wood warblers copied breeding‐site choices of poor quality conspecifics and despite temporary attraction to sites where the presence of good quality individuals was simulated, they did not ultimately settle near these individuals. Population density in the preceding year did not influence settlement patterns. We argue that when making settlement decisions, wood warblers assessed the expected level of local intraspecific competition and selectively copied breeding‐site choices of conspecifics or refused to settle, depending on competitive abilities of observed individuals. This adds a novel aspect to the patterns and processes of social information use proposed thus far, and provides support for the predicted negative effect of intraspecific competition on benefit of information. Moreover, it seems that habitat selection in wood warblers is a complex decision‐making process, in which initial decisions are adjusted after acquiring more accurate information. Synthesis Social information use in songbird habitat selection commonly involves copying the breeding‐site choices of conspecifics (so‐called conspecific attraction). To be adaptive, this strategy has to be discriminative, but almost no empirical studies have tested this assertion. Our study shows that birds may selectively use social information by copying settlement decisions of poor quality conspecifics, but avoid settling near good quality individuals, likely because of their high competitive abilities. This decision‐making pattern supports the predicted, yet not experimentally tested, tradeoff between information value and cost of competition in social information use. Our study highlights also that the use of social cues in settlement decisions may be both positively and negatively biased.     

Predation-Related Costs and Benefits of Conspecific Attraction in Songbirds—An Agent-Based Approach

Songbirds that follow a conspecific attraction strategy in the habitat selection process prefer to settle in habitat patches already occupied by other individuals. This largely affects the patterns of their spatio-temporal distribution and leads to clustered breeding. Although making informed settlement decisions is expected to be beneficial for individuals, such territory clusters may potentially provide additional fitness benefits (e.g., through the dilution effect) or costs (e.g., possibly facilitating nest localization if predators respond functionally to prey distribution). Thus, we hypothesized that the fitness consequences of following a conspecific attraction strategy may largely depend on the composition of the predator community. We developed an agent-based model in which we simulated the settling behavior of birds that use a conspecific attraction strategy and breed in a multi-predator landscape with predators that exhibited different foraging strategies. Moreover, we investigated whether Bayesian updating of prior settlement decisions according to the perceived predation risk may improve the fitness of birds that rely on conspecific cues. Our results provide evidence that the fitness consequences of conspecific attraction are predation-related. We found that in landscapes dominated by predators able to respond functionally to prey distribution, clustered breeding led to fitness costs. However, this cost could be reduced if birds performed Bayesian updating of prior settlement decisions and perceived nesting with too many neighbors as a threat. Our results did not support the hypothesis that in landscapes dominated by incidental predators, clustered breeding as a byproduct of conspecific attraction provides fitness benefits through the dilution effect. We suggest that this may be due to the spatial scale of songbirds’ aggregative behavior. In general, we provide evidence that when considering the fitness consequences of conspecific attraction for songbirds, one should expect a trade-off between the benefits of making informed decisions and the costs of clustering.     

Conspecific attraction and nest site selection in a nomadic species,the zebra finch

Conspecific nesting density affects many aspects of breeding biology, as well as habitat selection decisions. However, the large variations in breeding density observed in many species are yet to be fully explained. Here, we investigated the settlement patterns in a colonial species with variable breeding density and where resource distribution could be manipulated. The zebra finch, Taeniopygia guttata, is a classic avian model in evolutionary biology but we know surprisingly very little about nest site selection strategies and nesting densities in this species, and in fact, in nomadic species in general. Yet, important determinants of habitat selection strategies, including temporal predictability and breeding synchrony, are likely to be different in nomadic species than in the non‐nomadic species studied to date. Here, we manipulated the distribution of nesting sites (by providing nest boxes) and food patches (feeders) to test four non‐exclusive habitat selection hypotheses that could lead to nest aggregation: 1) attraction to resources, 2) attraction to breeding conspecifics, and 3) attraction to successful conspecifics and 4) use of private information (i.e. own reproductive success on a site). We found that wild zebra finches used conspecific presence and possibly reproductive success, to make decisions over where to locate their nests, but did not aggregate around water or food within the study areas. Moreover, there was a high degree of inter‐individual variation in nesting density preference. We discuss the significance of our results for habitat selection strategy in nomadic species and with respect to the differential selection pressures that individuals breeding at different densities may experience.     

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985–1990, we collected observational data in 1991–1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9–8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks. Received: 28 October 1996 / Accepted: 13 January 1997     

Testing the conspecific attraction hypothesis with Dupont's Larks,a resident species of songbird in central Spain

Some birds use social cues, such as the presence of conspecifics, when selecting breeding habitat. This phenomenon, known as conspecific attraction, has been well‐documented in migratory species, but has not been assessed for resident species of birds. We used Dupont's Larks (Chersophilus duponti) as a model species to determine if conspecific attraction plays a role in habitat selection by resident species of birds. At our study site in Soria province in central Spain, we monitored two potential habitat patches and one managed site where management actions had provided apparently suitable habitat. At each site, we broadcast recordings of the songs and calls of male Dupont's Larks, and monitored their presence during the breeding season and dispersal period in 2018 using automated recorders and field surveys. No birds were attracted to our study sites. Our results suggest that management of patches of suitable habitat should occur close to areas (within 1 km) already occupied by Dupont's Larks to encourage natural colonization because, based on our results, playback of conspecific vocalizations may not attract the species to new breeding areas. However, additional studies are needed before drawing conclusions about the effectiveness of conspecific attraction for this and other resident species of birds.     

When to use public information for breeding habitat selection? The role of environmental predictability and density dependence

To select their future breeding site, individuals usually have to assess local quality by using environmental cues. One optimal cue may be ‘public information’, the local reproductive success of conspecifics in a breeding patch (patch reproductive success) because it integrates the effect of all environmental factors on breeding success. However, the quality of information conveyed by patch reproductive success is likely to depend on (1) environmental predictability and (2) interactions between individuals. We investigated how these two factors, ignored by previous models, affect the performance of individuals using patch reproductive success for breeding habitat selection compared with other information. We built a two-patch, game-theoretical model to compare the success of a strategy of breeding habitat selection based on patch reproductive success relative to four other strategies: (1) random patch choice; (2) philopatry; (3) choice based on the presence of conspecifics the previous year (conspecific attraction); and (4) choice based on intrinsic patch quality the previous year. The results illustrate how the efficiency of strategies in tracking variations in patch quality depend on environmental predictability and costs linked to density dependence, themselves linked to the dynamics of spatial aggregation of individuals. In particular, strategies based on measures of patch reproductive success perform the best for intermediate and high temporal predictability of patch quality, whereas philopatry and conspecific attraction then perform poorly. The ‘conspecific attraction’ strategy always coexists with other strategies by efficiently parasitizing the information they use. We discuss the implications of a better understanding of breeding habitat selection behaviours for evolutionary and conservation biology. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Habitat matching: Alternatives and implications to populations and communities

Summary I evaluate habitat matching rules based on ideal distribution models of density-dependent habitat use. Recent approaches and the ideal free continuous input matching rule on which they depend, are restricted to only those habitats that are jointly occupied across the full range of population sizes. These assumptions may often be inappropriate to field applications of habitat matching. I develop alternatives that can be applied to a wide array of ideal forms of habitat selection, including the ideal free, continuous input example. Input matching can be distinguished from assumptions of consumer-resource models and preemptive habitat use by regressions of density between paired habitats (isodars). Isodars for continuous input models should be linear on a logarithmic scale, while those for consumer-resource models should be linear on an arithmetic scale. Pre-emptive isodars can be distinguished from the others by dramatic non-linearities at both low and high densities. Field data on white-footed mice support the consumer-resource theory. Implications of the rules for population regulation and community organization are highlighted by new models that specify how the fitness of pre-emptive habitat selectors should decline with increasing density. Strong non-linearities produced by comparisons between variable and homogeneous habitats produce reversing source-sink population regulation and a new form of cyclical community dynamics. Variable habitats act as a source of emigrants at low density and a sink for immigrants at high density. Subordinate species may occupy only the variable habitat at both low and high density.     

The effect of conspecific attraction on metapopulation dynamics

Random dispersal direction is assumed in all current metapopulation models. This assumption is called into question by recent experiments demonstrating that some species disperse preferentially to sites occupied by conspecifies. We incorporate conspecific attraction into two metapopulation models which differ in type of dispersal, the Levins model and a two-dimensional stepping-stone model. In both models, conspecific attraction lowers the proportion of occupied habitat patches within a metapopulation at equilibrium.     

Conspecific attraction in a grassland bird, the Baird's Sparrow

ABSTRACT.   Territorial songbirds generally use song to defend territories and attract mates, but conspecific song may also serve as a cue to attract other male songbirds to a breeding site. Although known to occur in some colonial and forest-associated species, only recently have investigators examined conspecific attraction in grassland species. We used a playback experiment to examine the possible role of conspecific attraction for males searching for potentially suitable breeding habitat in a grassland specialist, the Baird's Sparrow ( Ammodramus bairdii ). Experimental playback plots and control plots with similar landscape and vegetation characteristics were established at two sites in North Dakota. Baird's Sparrows colonized three of six experimental plots and none of six control plots. Males on experimental plots established territories adjacent to the playback stations and were sometimes observed counter-singing with the playback of conspecific songs. Vegetation characteristics were similar on all study plots, and did not explain differences in bird density on our treatment plots. Although we found that playback of conspecific songs attracted male Baird's Sparrows to previously unoccupied, potentially suitable habitat, further experiments are needed to examine the importance of conspecific attraction relative to other cues that birds may use, such as vegetation features. The conservation and management implications of conspecific attraction are not completely understood, but the presence of conspecifics should be considered as a potential cue in habitat selection by all species of birds.     

The heterospecific habitat copying hypothesis: can competitors indicate habitat quality?

According to the "habitat copying" hypothesis, animals use thereproductive performance of conspecifics to assess habitat suitabilityand choose their future breeding site. This is because conspecificsshare ecological needs and thus indicate habitat suitability.Here, we propose the "heterospecific habitat copying" hypothesis,which states that animals should use public information (i.e.,information derived from the performance of others) from con-and heterospecifics sharing ecological needs. In a correlationalapproach we test some assumptions and predictions of this hypothesiswith a data set from two sympatric bird populations, rollers(Coracias garrulus) and kestrels (Falco tinnunculus), usingthe same nest-boxes and exploiting similar food resources. Sincekestrels are residents and breed earlier, we assumed that theyare dominant over rollers for nest-box acquisition. The environmentappears to be patchy for both species and temporally predictablefor kestrels only. Two results suggest that the use of heterospecificpublic information in breeding habitat selection may be at work:(1) an increase in the reoccupancy probability by kestrels ofprevious roller nests with increasing nest success, and (2)an increase in roller breeding population with increasing localkestrel success. Most of the other observed patterns could beexplained by alternative mechanisms such as natal philopatry,breeding fidelity, conspecific attraction, intraspecific habitatcopying, and the effect of interspecific competition.     

Optimal floating and queuing strategies: consequences for density dependence and habitat loss

Field studies of many vertebrates show that some individuals (floaters) do not defend territories even when there is space for them to do so. We show that the evolutionarily stable strategy (ESS) for the threshold territory quality at which floating takes place is that which maximizes the size of the floating population (but not the total population, breeding population, or reproductive output). The ESS is solved separately for two assumptions: whether individuals wait to occupy a single territory or multiple territories and whether queuing rules are strict or if all waiting individuals are equally likely to obtain the next territory. The four combinations of these assumptions all give the same evolutionarily stable population size of both floaters and breeders. At the ESS, only territories with expected lifetime reproductive success (LRS) exceeding 1 should be occupied, which introduces a limit to ideal habitat selection. The behavioral decision to float alters the shape of the density-dependent response, reduces the equilibrium population size, and affects the response of the population to habitat loss. Specifically, the floater: breeder ratio is directly related to average breeding habitat quality, and the floater population size will decrease more than the breeding population size if better than average quality habitat is lost.     

Breeding habitat selection behaviors in heterogeneous environments: implications for modeling reintroduction

Animal movement and habitat selection behavior are important considerations in ecology, and remain a major issue for successful animal reintroductions. However, simple rules are often used to model movement or focus only on intrinsic environmental cues, neglecting recent insights in behavioral ecology on habitat selection processes. In particular, social information has been proposed as a widespread source of information for habitat evaluation.
We investigated the role of explicit breeding habitat selection strategies on the establishment pattern of reintroduced populations and their persistence. We considered local movement at the scale of a single population. We constructed a spatially-implicit demographic model that considered five breeding habitat selection rules: 1) random, 2) intrinsic habitat quality, 3) avoidance of conspecifics, 4) presence of conspecifics and 5) reproductive success of conspecifics. The impact of breeding habitat selection was examined for different release methods under various levels of environmental heterogeneity levels, for both long and short-lived monogamous species.
When heterogeneity between intrinsic habitat patch qualities is high, the persistence of reintroduced populations strongly depends on habitat selection strategies. Strategies based on intrinsic quality and conspecific reproductive success lead to a lower reintroduction failure risk than random, conspecific presence or avoidance-based strategies. Conspecific presence or avoidance-based strategies may aggregate individuals in suboptimal habitats. The release of adults seems to be more efficient independent of habitat selection strategy.
We emphasize the crucial role of oriented habitat selection behavior and non ideal habitat selection in movement modeling, particularly for reintroduction.     

Application of a predictive model to detect long-term changes in nest-site selection in the Bearded Vulture Gypaetus barbatus: conservation in relation to territory shrinkage

In this study we examined long-term variation in the selection of nesting cliffs for the recovering population of the Bearded Vulture Gypaetus barbatus inhabiting the Spanish Pyrenees. We focussed on variables indicating a high probability of cliff occupancy as determined by a previously published model. Although the breeding population increased from 53 to 93 territories between 1991 and 2002, the breeding range expanded only slightly. New and old nesting cliffs had similar habitat features in relation to topography, altitude and degree of human influence, but the distance between occupied cliffs was reduced (from 11.1 to 8.9 km). Thus the probabilities of occupation predicted by the model were lower for newly colonized locales. Our study shows that territory compression may occur without serious modification of nesting habitat quality. These results may arise from the lack of strong territorial behaviour by Bearded Vultures and the availability of high-quality cliffs. The relatively low quality of sites in adjacent mountains may prevent the expansion of the breeding range, but conspecific attraction may also play a role. Our study confirms that monitoring changes in key variables important to habitat selection is useful in determining long-term trends in settlement patterns in heterogeneous environments. The results also suggest that the available nest-site selection model may accurately predict cliff occupancy by Bearded Vultures in those areas where the distance to the nearest neighbour is not a limiting factor. In particular, the model may be useful in establishing priority areas for reintroduction.     

Territorial aggression can be sensitive to the status of heterospecific intruders

Territorial animals are known to be able to differentiate between intruding individuals posing a low or high threat and adjust their aggressive response accordingly. However, plastic territorial aggression based on recognising individuals with different attributes is typically assumed to be relevant only in the context of conspecific interactions. In this study, we investigated territorial aggression of neotropical cichlid fish in their natural habitat to assess whether responses to different types of individuals of another species can also be plastic. We show that arrow cichlids (Amphilophus zaliosus) adjusted their territorial aggression regarding the status of heterospecific intruders: breeding individuals of Amphilophus astorquii received a lower level of aggression than non-breeders. The same pattern was also found for the two different types of A. astorquii individuals intruding into conspecific territories. These results suggest that heterospecific individuals should not be ignored when considering selection pressures shaping plasticity of aggressive behaviour in territorial animals.     

Distribution dynamics of a great bustard metapopulation throughout a decade: influence of conspecific attraction and recruitment

Dispersing individuals can use conspecifics as indicators of habitat quality and aggregate at traditionally occupied sites, leaving other favourable patches unoccupied. Here we test the predictions of the conspecific-based habitat selection hypothesis on a Spanish great bustard (Otis tarda) metapopulation, currently fragmented due to recent human-induced habitat changes. The number of birds had increased by 23% between 1988 and 1998, but not consistently among leks. Leks that were large in 1988 increased, while those that were small decreased, which suggests that dispersing individuals used the numbers of conspecifics as cues for breeding-site selection. Moreover, leks with high productivity increased, while those with low productivity decreased. Finally, lek distribution was markedly stable throughout the decade, with no establishment of new leks, and suitable habitat patches remained unoccupied, as predicted by the conspecific attraction hypothesis. These results were corroborated by a simulation model which incorporated natal dispersal rates between leks as obtained through radio-tracking of 15 birds that survived throughout their 4-year dispersal period. In conclusion, in spite of the apparent increase in total numbers throughout the decade, both conspecific attraction and local differences in reproductive success contributed to a more aggregated distribution, increasing the species' vulnerability to local catastrophes, and the risks of reduced genetic diversity and extinction of small leks.     

Managing breeding forest songbirds with conspecific song playbacks

Recent studies confirm that several territorial songbird species use conspecific cues, including song, when selecting habitat. We conducted a controlled experiment with a population of black-throated blue warblers Dendroica caerulescens , a species of concern in boreal hardwood transition forests, to determine whether song playbacks influence settlement patterns. We tested whether conspecific song broadcast during spring arrival attracts black-throated blue warblers during territory establishment, and whether song playbacks shift territories from the previous year's breeding distributions. The results of our experiment demonstrate that song playbacks significantly increased territorial occupancy and shifted territories closer to the speakers. Loss of breeding habitat is implicated in population declines of many songbirds, and our results suggest that song playbacks should be further investigated as a tool for mitigating the effects of disruptive management actions. Additional work should consider the spatial arrangement of playback speakers to avoid attracting birds from safe areas. The demographic consequences of conspecific attraction must also be explored before this technique is widely implemented.     

Emergent properties of conspecific attraction in fragmented landscapes

Attraction to conspecifics may have wide-ranging implications for habitat selection and metapopulation theory, yet little is known about the process of attraction and its effects relative to other habitat selection strategies. Using individual-based simulations, I investigated the emergent properties of conspecific attraction during habitat selection on survival, fecundity, short-term fitness (survival x fecundity), and distributions in fragmented landscapes. I simulated conspecific attraction during searching and settlement decisions and compared attraction with random, habitat-based (searching for the presence of habitat), and habitat quality sampling strategies (searching for and settling in high-quality habitat). Conspecific attraction during searching or settlement decisions had different consequences for animals: attraction while searching increased survival by decreasing time spent in nonsuitable habitat, whereas attraction during settlement increased fecundity by aggregating animals in high-quality habitats. Habitat-based sampling did not improve fitness over attraction, but directly sampling habitat quality resulted in the highest short-term fitness among strategies. These results suggest that attraction can improve fitness when animals cannot directly assess habitat quality. Interestingly, conspecific attraction influenced distributions by generating patch size effects and weak edge effects, highlighting that attraction is one potential, yet previously unappreciated, mechanism to explain the widespread patterns of animal sensitivity to habitat fragmentation.