Allee effects and conspecific cueing jointly lead to conspecific attraction

Conspecific attraction is the preferential settlement into habitat patches with conspecifics. To be a good proximate strategy, fitness gains from settling with conspecifics must outweigh the costs of higher conspecific densities, such as intraspecific competition. Two types of benefits have been proposed to explain conspecific attraction: Allee effects (i.e., positive density dependence) and conspecific cueing (using conspecifics as an indicator of habitat quality). I present empirical evidence for conspecific attraction in the settlement of the porcelain crab, Petrolisthes cinctipes Randall (Anomura: Porcellanidae). Previous work demonstrated that P. cinctipes experiences strong intraspecific competition and that both Allee effects and conspecific cueing are present in P. cinctipes life-history. I developed an empirically-based fitness model of the costs and benefits of settling with conspecifics. Based on this model, I simulated optimal settlement to habitat patches that varied in conspecific density and habitat quality, where the correlation between density and habitat quality determined the level of conspecific cueing. I tested whether Allee effects alone, conspecific cueing alone, or Allee effects and conspecific cueing together could provide an ultimate explanation for the proximate settlement behavior of P. cinctipes. The settlement simulation was consistent with empirical settlement only when Allee effects and conspecific cueing were both included. Three life-history features are critical to this conclusion: (1) fitness is maximized at intermediate density, (2) fitness depends on the decisions of previous settlers, and (3) conspecific density provides good information about habitat quality. The quality of information garnered from conspecifics determines whether conspecific attraction is a good proximate strategy for settlement. I present a graphical illustration demonstrating how Allee effects and conspecific cueing work together to influence fitness, providing a conceptual framework for other systems.Electronic Supplementary Material Supplementary material is available for this article at and is accessible for authorized users.     

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.     

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 is a Missing Component in Wildlife Habitat Modeling

Abstract Wildlife biologists use knowledge about wildlife-habitat relationships to create habitat models to predict species occurrence across a landscape. Researchers attribute limitations in predictive ability of a habitat model to data deficiencies, missing parameters, error introduced by specifications of the statistical model, and natural variation. Few wildlife biologists, however, have incorporated intra- and interspecific interactions (e.g., conspecific attraction, competition, predator-prey relationships) to increase predictive accuracy of habitat models. Based on our literature review and preliminary data analysis, conspecific attraction can be a primary factor influencing habitat selection in wildlife. Conspecific attraction can lead to clustered distributions of wildlife within available habitat, reducing the predictive ability of habitat models based on vegetative and geographic parameters alone. We suggest wildlife biologists consider incorporating a parameter in habitat models for the clustered distribution of individuals within available habitat and investigate the mechanisms leading to clustered distributions of species, especially conspecific attraction.     

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”.     

The nose knows: linking sensory cue use,settlement decisions,and post-settlement survival in a temperate reef fish

Habitat selection by animals that migrate or disperse ultimately determines the biotic and abiotic environment they will experience in subsequent life stages. Intuitively, for habitat selection to be adaptive, animals should respond positively to cues produced by habitat characteristics that will enhance their fitness in the new environment. However, there are many examples of dispersing animals where individuals are attracted to cues produced by factors that reduce their fitness after arrival. In this study, we use a temperate reef fish to examine the relative importance of habitat-associated cues in habitat selection decisions, and assess whether use of these cues is adaptive across early life stages. We used a series of laboratory- and field-based manipulative experiments to test: (1) what habitat-associated cues are likely used to locate suitable habitat; (2) whether in situ settlement patterns reflect the cue response tested in the laboratory; and (3) whether the aspects of the habitat that stimulate settlement are the same as those that maximize survival. We observed a positive response to multiple habitat-associated cues, with conspecific cues eliciting the strongest behavioral response in laboratory choice experiments, and a strong inverse density-dependent relationship at settlement. Macroalgal cues also elicited a positive response at settlement, but were associated with higher mortality after settlement, suggesting that habitat selection decisions are not always adaptive. We argue that this non-intuitive behavior may still be adaptive if it improves fitness at an earlier life stage, as habitat selection behavior is the result of tradeoffs in fitness costs across multiple stages.     

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.     

Linking Dynamic Habitat Selection with Wading Bird Foraging Distributions across Resource Gradients

Species distribution models (SDM) link species occurrence with a suite of environmental predictors and provide an estimate of habitat quality when the variable set captures the biological requirements of the species. SDMs are inherently more complex when they include components of a species’ ecology such as conspecific attraction and behavioral flexibility to exploit resources that vary across time and space. Wading birds are highly mobile, demonstrate flexible habitat selection, and respond quickly to changes in habitat quality; thus serving as important indicator species for wetland systems. We developed a spatio-temporal, multi-SDM framework using Great Egret (Ardea alba), White Ibis (Eudocimus albus), and Wood Stork (Mycteria Americana) distributions over a decadal gradient of environmental conditions to predict species-specific abundance across space and locations used on the landscape over time. In models of temporal dynamics, species demonstrated conditional preferences for resources based on resource levels linked to differing temporal scales. Wading bird abundance was highest when prey production from optimal periods of inundation was concentrated in shallow depths. Similar responses were observed in models predicting locations used over time, accounting for spatial autocorrelation. Species clustered in response to differing habitat conditions, indicating that social attraction can co-vary with foraging strategy, water-level changes, and habitat quality. This modeling framework can be applied to evaluate the multi-annual resource pulses occurring in real-time, climate change scenarios, or restorative hydrological regimes by tracking changing seasonal and annual distribution and abundance of high quality foraging patches.     

The Effect of Adult Aggression on Habitat Selection by Settlers of Two Coral-Dwelling Damselfishes

Coral-reef fishes experience a major challenge when facing settlement in a multi-threat environment, within which, using settlement cues, they need to select a suitable site. Studies in laboratories and artificial setups have shown that the presence of conspecific adults often serves as a positive settlement cue, whose value is explained by the increased survival of juveniles in an already proven fit environment. However, settlement in already inhabited corals may expose the recruits to adult aggression. Daily observations and manipulation experiments were used in the present study, which was conducted in the natural reef. We revealed differential strategies of settlers, which do not necessarily join conspecific adults. Dascyllus aruanus prefer to settle near (not with) their aggressive adults, and to join them only after gaining in size; whereas Dascyllus marginatus settlers in densely populated reefs settle independently of their adult distribution. Our results present different solutions to the challenges faced by fish recruits while selecting their microhabitat, and emphasize the complexity of habitat selection by the naïve settlers. Although laboratory experiments are important to the understanding of fish habitat selection, further studies in natural habitats are essential in order to elucidate the actual patterns of settlement and habitat selection, which are crucial for the survival of coral-reef fish populations.     

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.     

Multi-scale habitat selection affects offspring survival in a precocial species

In theory, habitat preferences should be adaptive. Accordingly, fitness is often assumed to be greater in preferred habitats; however, this assumption is rarely tested and, when it is, the results are often equivocal. Habitat preferences may not directly convey fitness advantages if animals are constrained by tradeoffs with other selective pressures like predation or food availability. We address unresolved questions about the survival consequences of habitat choices made during brood-rearing in a precocial species with exclusive maternal care (mallard Anas platyrhynchos, n = 582 radio-marked females on 27 sites over 8 years). We directly linked duckling survival with habitat selection patterns at two spatial scales using logistic regression and model selection techniques. At the landscape scale (55–80 km²), females that demonstrated stronger selection of areas with more cover type 4 wetlands and greater total cover type 3 wetland area (wetlands with large expanses of open water surrounded by either a narrow or wide peripheral band of vegetation, respectively) had lower duckling survival rates than did females that demonstrated weaker selection of these habitats. At finer scales (0.32–7.16 km²), females selected brood-rearing areas with a greater proportion of wetland habitat with no consequences for duckling survival. However, females that avoided woody perennial habitats composed of trees and shrubs fledged more ducklings. The relationship between habitat selection and survival depended on both spatial scale and habitats considered. Females did not consistently select brood-rearing habitats that conferred the greatest benefits, an unexpected finding, although one that has also been reported in other recent studies of breeding birds.     

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.     

Selective sweeps for recessive alleles and for other modes of dominance

A selective sweep describes the reduction of linked genetic variation due to strong positive selection. If s is the fitness advantage of a homozygote for the beneficial allele and h its dominance coefficient, it is usually assumed that h = 1/2, i.e. the beneficial allele is co-dominant. We complement existing theory for selective sweeps by assuming that h is any value in [0, 1]. We show that genetic diversity patterns under selective sweeps with strength s and dominance 0 < h < 1 are similar to co-dominant sweeps with selection strength 2hs. Moreover, we focus on the case h = 0 of a completely recessive beneficial allele. We find that the length of the sweep, i.e. the time from occurrence until fixation of the beneficial allele, is of the order of \({\sqrt{N/s}}\) generations, if N is the population size. Simulations as well as our results show that genetic diversity patterns in the recessive case h = 0 greatly differ from all other cases.     

A review of conspecific attraction for habitat selection across taxa

Many species across taxa select habitat based on conspecific presence, known as conspecific attraction. Studies that document conspecific attraction typically provide social information (i.e., cues that indicate the presence of a given species) and then determine if a given species is more likely to settle at locations where the social information is provided compared to those locations that do not. Although the number of studies examining conspecific attraction has grown in recent years, a comprehensive review has not yet been undertaken. Here, we conducted a review of the literature and found 151 studies investigating conspecific attraction across eight taxa. We found that conspecific attraction is widespread with between 80% and 100% of studies, depending on taxa, documenting positive associations between habitat selection and the presence of conspecific cues. Conspecific attraction has been documented more frequently in bird and fish species with less attention given to invertebrate and mammal species. We use the patterns we found to (a) provide an overview of the current state of research on conspecific attraction and (b) discuss how important factors, such as cue characteristics and life history traits, may play a role in shaping conspecific attraction patterns within and across taxa.     

Phenotypic and transgenerational plasticity promote local adaptation to sun and shade environments

Adaptive plasticity is expected to be important when the grain of environmental variation is encompassed in offspring dispersal distance. We investigated patterns of local adaptation, selection and plasticity in an association of plant morphology with fine-scale habitat shifts from oak canopy understory to adjacent grassland habitat in Claytonia perfoliata. Populations from beneath the canopy of oak trees were >90 % broad leaved and large seeded, while plants from adjacent grassland habitat were >90 % linear-leaved and small seeded. In a 2-year study, we used reciprocal transplants and phenotypic selection analysis to investigate local adaptation, selection, plasticity and maternal effects in this trait-environment association. Transgenerational effects were studied by planting offspring of inbred maternal families grown in both environments across the same environments in the second year. Reciprocal transplants revealed local adaptation to habitat type: broad-leaved forms had higher fitness in oak understory and linear-leaved plants had higher fitness in open grassland habitat. Phenotypic selection analyses indicated selection for narrower leaves and lower SLA in open habitat, and selection for broad leaves and intermediate values of SLA in understory. Both plant morphs exhibited plastic responses in traits in the same direction as selection on traits (narrower leaves and lower SLA in open habitat) suggesting that plasticity is adaptive. We detected an adaptive transgenerational effect in which maternal environment influenced offspring fitness; offspring of grassland-reared plants had higher fitness than understory-reared plants when grown in grassland. We did not detect costs of plasticity, but did find a positive association between leaf shape plasticity and fitness in linear-leaved plants in grassland habitat. Together, these findings indicate that fixed differences in trait values corresponding to selection across habitat contribute to local adaptation, but that plasticity and maternal environmental effects may be favored through promotion of survival across heterogeneous environments.     

The effects of conspecifics on burrow selection in juvenile spotted salamanders (<Emphasis Type="Italic">Ambystoma maculatum)</Emphasis>

Spotted salamanders (Ambystoma maculatum) are pond-breeding amphibians that disperse into terrestrial habitat from natal wetlands after undergoing metamorphosis, relying on small-mammal burrows and coarse woody debris for refugia. The effect of conspecifics on burrow use in juvenile salamanders is poorly understood. Determining how the presence of conspecifics influences the settlement decisions of juvenile salamanders can increase our understanding of amphibian dispersal and our ability to predict population dynamics. We conducted behavioral laboratory trials using 58 recently metamorphosed salamanders to examine how salamanders selected burrows in the presence of conspecifics. Salamanders were more likely to settle in a burrow that was occupied by a conspecific versus an unoccupied burrow. Our results indicate that juvenile salamanders may show conspecific attraction and/or trailing behavior during the dispersal phase.     

Nonideal nest box selection by tree swallows breeding in farmlands: Evidence for an ecological trap?

Animals are expected to select a breeding habitat using cues that should reflect, directly or not, the fitness outcome of the different habitat options. However, human‐induced environmental changes can alter the relationships between habitat characteristics and their fitness consequences, leading to maladaptive habitat choices. The most severe case of such nonideal habitat selection is the ecological trap, which occurs when individuals prefer to settle in poor‐quality habitats while better ones are available. Here, we studied the adaptiveness of nest box selection in a tree swallow (Tachycineta bicolor) population breeding over a 10‐year period in a network of 400 nest boxes distributed along a gradient of agricultural intensification in southern Québec, Canada. We first examined the effects of multiple environmental and social habitat characteristics on nest box preference to identify potential settlement cues. We then assessed the links between those cues and habitat quality as defined by the reproductive performance of individuals that settled early or late in nest boxes. We found that tree swallows preferred nesting in open habitats with high cover of perennial forage crops, high spring insect biomass, and high density of house sparrows (Passer domesticus), their main competitors for nest sites. They also preferred nesting where the density of breeders and their mean number of fledglings during the previous year were high. However, we detected mismatches between preference and habitat quality for several environmental variables. The density of competitors and conspecific social information showed severe mismatches, as their relationships to preference and breeding success went in opposite direction under certain circumstances. Spring food availability and agricultural landscape context, while related to preferences, were not related to breeding success. Overall, our study emphasizes the complexity of habitat selection behavior and provides evidence that multiple mechanisms may potentially lead to an ecological trap in farmlands.     

Evolutionary determinants of population differences in population growth rate × habitat temperature interactions in Chironomus riparius

Little is known about intraspecific variation in fitness performance in response to thermal stress among natural populations and how this relates to evolutionary aspects of species ecology. In this study, population growth rate (PGR; a composite fitness measure) varied among five natural Chironomus riparius populations sampled across a climatic gradient when subjected to three temperature treatments reflecting the typical range of summer habitat temperatures (20, 24 and 28 °C). The variation could be explained by a complex model including effects of genetic drift, genetic diversity and adaptation to average temperature during the warmest month, in addition to experimental temperature. All populations suffered a decrease in PGR from 20 to 28 °C and ΔPGR was significantly correlated with the respective average habitat temperature in the warmest month—populations from warmer areas showing lower ΔPGR. This implies that long-term exposure to higher temperatures in the warmest month (the key reproductive period for C. riparius) is likely to be a key selective force influencing fitness at higher temperatures. A comparison of phenotypic divergence and neutral genetic differentiation revealed that one phenotypic trait—the number of fertile egg masses per female—appeared to be under positive selection in some populations. Our findings support a role for response to temperature selection along a climatic gradient and suggest population history is a key determinant of intraspecific fitness variation. We stress the importance of integrating different types of data (climatic, experimental, genetic) in order to understand the effects of global climate change on biodiversity.     

Disentangling drivers of spatial autocorrelation in species distribution models

Species distribution models (SDMs) are frequently used to understand the influence of site properties on species occurrence. For robust model inference, SDMs need to account for the spatial autocorrelation of virtually all species occurrence data. Current methods do not routinely distinguish between extrinsic and intrinsic drivers of spatial autocorrelation, although these may have different implications for conservation. Here, we present and test a method that disentangles extrinsic and intrinsic drivers of spatial autocorrelation using repeated observations of a species. We focus on unknown habitat characteristics and conspecific interactions as extrinsic and intrinsic drivers, respectively. We model the former with spatially correlated random effects and the latter with an autocovariate, such that the spatially correlated random effects are constant across the repeated observations whereas the autocovariate may change. We tested the performance of our model on virtual species data and applied it to observations of the corncrake Crex crex in the Netherlands. Applying our model to virtual species data revealed that it was well able to distinguish between the two different drivers of spatial autocorrelation, outperforming models with no or a single component for spatial autocorrelation. This finding was independent of the direction of the conspecific interactions (i.e. conspecific attraction versus competitive exclusion). The simulations confirmed that the ability of our model to disentangle both drivers of autocorrelation depends on repeated observations. In the case study, we discovered that the corncrake has a stronger response to habitat characteristics compared to a model that did not include spatially correlated random effects, whereas conspecific interactions appeared to be less important. This implies that future conservation efforts should primarily focus on maximizing habitat availability. Our study shows how to systematically disentangle extrinsic and intrinsic drivers of spatial autocorrelation. The method we propose can help to correctly identify the main drivers of species distributions.     

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.