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.     

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.     

Habitat selection rules in breeding mallards (Anas platyrhynchos): a test of two competing hypotheses

Ideal preemption and conspecific attraction are alternative hypotheses of the habitat selection rules used by individuals. According to the former an occupied site is assumed to be preempted and therefore not available for later arriving individuals, whereas according to the latter individuals are assumed to be attracted by conspecifics to occupied sites, rather than avoiding them. We studied these competing hypotheses in breeding mallards (Anas platyrhynchos) by a cross-over experiment in 2 years, introducing birds onto lakes before migratory wild mallards arrived. If mallards use the ideal preemptive rule, breeding density of wild mallards in experimental lakes should be lower and they should be occupied less frequently than control lakes, but if mallards use the conspecific attraction rule the reverse should be true. Our results allowed us to reject the ideal preemptive rule whereas the conspecific attraction rule was to some extent supported. We discuss these findings in relation to population limitation. The results suggest that the local breeding population studied is not limited by spacing behaviour related to habitat selection. Received: 14 July 1997 / Accepted: 17 November 1997     

Does attraction to conspecifics explain the patch‐size effect? An experimental test

Recent theory suggests that attraction to conspecifics during habitat selection can be one potential, yet untested, mechanism for animal sensitivity to habitat fragmentation. The least flycatcher Empidonax minimus , a highly territorial migratory bird, has previously been shown to be attracted to conspecifics and sensitive to patch size by avoiding small patches of riparian forest in Montana, USA. I used a large-scale field experiment in this region to test the conspecific attraction hypothesis for explaining sensitivity to patch size, and I supplemented this experiment by estimating whether vegetation structure, nest predation, or nest parasitism rates could better explain patterns of sensitivity to patch size. Vegetation structure did not vary consistently with patch size, based on a random sample of patches across 150  km of the Madison and Missouri Rivers, Montana. Nest predation and parasitism rates by brown-headed cowbirds Molothrus ater also did not vary with patch size during the experiment. However, when conspecific cues were simulated across a gradient of patch sizes, flycatchers settled in all patches – and their sensitivity to patch size vanished – providing strong support for the conspecific attraction hypothesis. These results provide the first experimental evidence that attraction to conspecifics can indeed help explain area sensitivity in nature and highlight how understanding the role of animal behavior in heterogeneous landscapes can aid in interpreting pressing conservation issues.     

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.     

Migration of the clouded Apollo butterfly Parnassius mnemosyne in a network of suitable habitats – effects of patch characteristics

The clouded Apollo Parnassius mnemosyne is a food plant specialist with short but frequent movements between habitat patches. The short average dispersal distances suggest that the probability of colonisation of vacant patches decreases rapidly as the distance between the source and target patches increases, which means that a dense habitat network is needed for the conservation of the species. Both emigration rate and the number of immigrants varied among patches and were not affected only by isolation but also by several other patch characteristics. The model that explained most of the variation in emigration rates among patches included patch area and the number of conspecifics. The area and the population density of the target patch had significant effects on the number of arriving immigrants. Thus, the colonisation of vacant patches is dependent on these patch characteristics. Generally, emigration rates were lower and residence times longer in large patches with many conspecifics. Butterfly density was the most important single factor explaining the variation in the number of immigrants among patches, although the positive effect of the area of the target patch was also significant. As a consequence of the marked positive density dependence caused by conspecific attraction, small patches with higher than average butterfly density, receive more immigrants than could be expected based on the patch area only. Due to conspecific attraction, per capita immigration rates are higher in small than large patches. Thus, immigration may have a more significant effect on the local dynamics of small than large populations.     

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.     

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.     

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.     

Space use and social association in a gregarious ungulate: Testing the conspecific attraction and resource dispersion hypotheses

Animals use a variety of proximate cues to assess habitat quality when resources vary spatiotemporally. Two nonmutually exclusive strategies to assess habitat quality involve either direct assessment of landscape features or observation of social cues from conspecifics as a form of information transfer about forage resources. The conspecific attraction hypothesis proposes that individual space use is dependent on the distribution of conspecifics rather than the location of resource patches, whereas the resource dispersion hypothesis proposes that individual space use and social association are driven by the abundance and distribution of resources. We tested the conspecific attraction and the resource dispersion hypotheses as two nonmutually exclusive hypotheses explaining social association and of adult female caribou (Rangifer tarandus). We used location data from GPS collars to estimate interannual site fidelity and networks representing home range overlap and social associations among individual caribou. We found that home range overlap and social associations were correlated with resource distribution in summer and conspecific attraction in winter. In summer, when resources were distributed relatively homogeneously, interannual site fidelity was high and home range overlap and social associations were low. Conversely, in winter when resources were distributed relatively heterogeneously, interannual site fidelity was low and home range overlap and social associations were high. As access to resources changes across seasons, caribou appear to alter social behavior and space use. In summer, caribou may use cues associated with the distribution of forage, and in winter caribou may use cues from conspecifics to access forage. Our results have broad implications for our understanding of caribou socioecology, suggesting that caribou use season‐specific strategies to locate forage. Caribou populations continue to decline globally, and our finding that conspecific attraction is likely related to access to forage suggests that further fragmentation of caribou habitat could limit social association among caribou, particularly in winter when access to resources may be limited.     

Spatio‐temporal dynamics of density‐dependent dispersal during a population colonisation

Predicting population colonisations requires understanding how spatio‐temporal changes in density affect dispersal. Density can inform on fitness prospects, acting as a cue for either habitat quality, or competition over resources. However, when escaping competition, high local density should only increase emigration if lower‐density patches are available elsewhere. Few empirical studies on dispersal have considered the effects of density at the local and landscape scale simultaneously. To explore this, we analyze 5 years of individual‐based data from an experimental introduction of wild guppies Poecilia reticulata. Natal dispersal showed a decrease in local density dependence as density at the landscape level increased. Landscape density did not affect dispersal among adults, but local density‐dependent dispersal switched from negative (conspecific attraction) to positive (conspecific avoidance), as the colonisation progressed. This study demonstrates that densities at various scales interact to determine dispersal, and suggests that dispersal trade‐offs differ across life stages.     

Finding a place to live: conspecific attraction affects habitat selection in juvenile green and golden bell frogs

Conspecific attraction plays an important role in habitat selection of several taxa and can affect and determine distribution patterns of populations. The behaviour is largely studied and widespread among birds, but in amphibians, its occurrence seems limited to breeding habitats of adults and gregarious tadpoles. The Australian green and golden bell frogs (Litoria aurea) have suffered considerable shrinking of their original distribution in south-eastern Australia since the 1970s. Currently, with only about 40 populations remaining, the species is considered nationally threatened. In natural conditions, these frogs are aggregated in the landscape and do not seem to occupy all suitable ponds within the occurrence area. To date, studies focusing on the frogs’ habitat have failed in finding a general habitat feature that explains current or past occupancy. This led us to the hypothesis that social cues may play a key role in habitat selection in this species. Using two choice experiments, we tested the preference of juvenile green and golden bell frogs for habitats containing cues of conspecifics of similar size versus habitats without conspecific cues. Tested frogs did not show a preference for habitats containing only scent from conspecifics but did prefer habitats where conspecifics were present. Our results show that conspecific attraction is a determining factor in juvenile green and golden bell frog habitat selection. To our knowledge, this is the first time the behaviour is shown to occur in juvenile frogs in the habitat selection context. From a conservation management point of view, the behaviour may help to explain the failure of reintroductions to areas where the frogs have been extinct, and the non-occupation of suitable created habitats in areas where they still inhabit and develop appropriated management strategies.     

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.     

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.     

Social information in nest colonisation and occupancy in a long-lived, solitary breeding bird

Recent work increasingly reveals the importance of social information in individual dispersal decisions, population dynamics and conservation. Much of the knowledge gained to date comes from studies on short-lived and/or densely breeding species. In contrast, our understanding of the processes involved in nest-site selection for long-lived, solitary breeding species is insufficient. We increased nest-site availability by nest-box supplementation over a 5-year period in a population of a long-lived, solitary, secondary-cavity nesting bird, the European roller Coracias garrulus, breeding in natural cavities and human constructions. We tested the nest limitation and the inadvertent conspecific social information hypothesis in order to study the dynamics and mechanisms of abandonment of previously used nests and the colonisation of new ones. Our data lend support to the nest-limitation hypothesis both in terms of quantity-population and the size of breeding clusters increased, and suitability--the majority of pairs used and re-occupied nest-boxes. Nevertheless, the use of natural cavities did not decrease after 5 years. At the between-patch scale, rollers were revealed to colonise nest-boxes based on conspecific social attraction, namely distance to the nearest neighbour in the same season. Despite the unpredictability of patch productivity, at the within--patch scale, the selection of previously unoccupied cavities was consistent with the performance-based conspecific attraction hypothesis. Philopatry could account for the repeated use of cavities, because nests that were used for two successive years were more likely to also be reused in the subsequent season.     

Public information affects breeding dispersal in a colonial bird: kittiwakes cue on neighbours

Habitat selection and dispersal behaviour are key processes in evolutionary ecology. Recent studies have suggested that individuals may use the reproductive performance of conspecifics as a source of public information on breeding patch quality for dispersal decisions, but experimental evidence is still limited for species breeding in aggregates, i.e. colonial species. We addressed this issue by manipulating the local breeding success of marked individuals and that of their neighbours on a series of breeding patches of a colonial seabird, the black-legged kittiwake (Rissa tridactyla). Based on previous observations in this species, we predicted that individuals that lost their eggs on successful patches would attend their nest and come back to it the year after at a higher rate than individuals that lost their eggs on patches where their neighbours were also in failure. As predicted, the attendance of breeders and prospectors was strongly affected by the local level of breeding success, resulting in differential site fidelity and recruitment. This suggests that individuals used information conveyed by conspecific breeding performance to make decisions relative to breeding site selection. This process can amplify the response of these populations to environmental change and may have contributed to the evolution of colonial breeding.     

Climate and density influence annual survival and movement in a migratory songbird

Assessing the drivers of survival across the annual cycle is important for understanding when and how population limitation occurs in migratory animals. Density‐dependent population regulation can occur during breeding and nonbreeding periods, and large‐scale climate cycles can also affect survival throughout the annual cycle via their effects on local weather and vegetation productivity. Most studies of survival use mark–recapture techniques to estimate apparent survival, but true survival rates remain obscured due to unknown rates of permanent emigration. This is especially problematic when assessing annual survival of migratory birds, whose movement between breeding attempts, or breeding dispersal, can be substantial. We used a multistate approach to examine drivers of annual survival and one component of breeding dispersal (habitat‐specific movements) in a population of American redstarts (Setophaga ruticilla) over 11 years in two adjacent habitat types. Annual survival displayed a curvilinear relation to the Southern Oscillation Index, with lower survival during La Niña and El Niño conditions. Although redstart density had no impact on survival, habitat‐specific density influenced local movements between habitat types, with redstarts being less likely to disperse from their previous year's breeding habitat as density within that habitat increased. This finding was strongest in males and may be explained by conspecific attraction influencing settlement decisions. Survival was lowest in young males, but movement was highest in this group, indicating that apparent survival rates were likely biased low due to permanent emigration. Our findings demonstrate the utility of examining breeding dispersal in mark–recapture studies and complement recent work using spatially explicit models of dispersal probability to obtain greater accuracy in survival estimates.     

Spatial patterns of the frog Oophaga pumilio in a plantation system are consistent with conspecific attraction

The conspecific attraction hypothesis predicts that individuals are attracted to conspecifics because conspecifics may be cues to quality habitat and/or colonists may benefit from living in aggregations. Poison frogs (Dendrobatidae) are aposematic, territorial, and visually oriented—three characteristics which make dendrobatids an appropriate model to test for conspecific attraction. In this study, we tested this hypothesis using an extensive mark‐recapture dataset of the strawberry poison frog (Oophaga pumilio) from La Selva Biological Station, Costa Rica. Data were collected from replicate populations in a relatively homogenous Theobroma cacao plantation, which provided a unique opportunity to test how conspecifics influence the spatial ecology of migrants in a controlled habitat with homogenous structure. We predicted that (1) individuals entering a population would aggregate with resident adults, (2) migrants would share sites with residents at a greater frequency than expected by chance, and (3) migrant home ranges would have shorter nearest‐neighbor distances (NND) to residents than expected by chance. The results were consistent with these three predictions: Relative to random simulations, we observed significant aggregation, home‐range overlap, and NND distribution functions in four, five, and six, respectively, of the six migrant–resident groups analyzed. Conspecific attraction may benefit migrant O. pumilio by providing cues to suitable home sites and/or increasing the potential for social interactions with conspecifics; if true, these benefits should outweigh the negative effects of other factors associated with aggregation. The observed aggregation between migrant and resident O. pumilio is consistent with conspecific attraction in dendrobatid frogs, and our study provides rare support from a field setting that conspecific attraction may be a relevant mechanism for models of anuran spatial ecology.     

Behavioural mechanisms that undermine species envelope models: the causes of patchiness in the distribution of great bustards Otis tarda in Spain

Despite the general success of species envelope models, capturing the fine-scale detail of patchiness in the distributions of some species is problematic. For great bustards in Spain, apparently suitable habitat patches remain unoccupied and cannot be distinguished from occupied patches in current distribution models. We consider philopatry and conspecific attraction as main behavioural mechanisms which could account for this patchiness, and then look for evidence of their influence on the distribution of great bustards across the whole of Spain. We compared the characteristics of habitat patches classed as suitable by a distribution model according to whether they were actually used or not. Occupied patches were larger than unoccupied patches and over-used in proportion to their size, suggesting aggregation and a metapopulation structure. Arguing that conspecific attraction may serve to transfer information about site history and environmental predictability (at least over a short time period), we compared the coefficients of variation in time-series of vegetation and climatic factors at occupied and unoccupied sites. Great bustards chose sites which were more environmentally stable at critical periods in the breeding cycle, "public information" that can only be gained from others rather than through sampling. There is thus evidence that both metapopulation dynamics and conspecific attraction influence the large scale distribution of great bustards in Spain. We discuss how alternative predictor variables and multi-stage analyses may help us to incorporate behavioural mechanisms into distribution models, but acknowledge that there are limits to the value of species envelope models for animal species making decisions.     

Inter-individual variability and conspecific densities: consequences for population regulation and range expansion

The presence of conspecifics can strongly modulate the quality of a breeding site. Both positive and negative effects of conspecifics can act on the same individuals, with the final balance between its costs and benefits depending on individual characteristics. A particular case of inter-individual variation found in many avian species is chromatic variability. Among birds, plumage coloration can co-vary with morphology, physiology and behavior as well as with age. These relationships suggest that cost-benefit balances of conspecific presence may be different for individuals with different colorations. We investigated whether inter-individual variability affects population regulation and expansion processes by analyzing potential differences in density-dependent productivity and settlement patterns in relation to plumage coloration in a population of a long-lived avian species recently undergoing a notable increase in numbers and distribution range. Our results show strong variation in the effect of density on productivity of breeding pairs depending on plumage coloration of their members. Productivity of dark birds decreased along the breeding density gradient while that of lighter breeders remained unchanged with conspecific density. In a similar way, our results showed an uneven occupation of localities by individuals with different plumage coloration in relation to local densities, with the breeding of lighter harriers more aggregated than that of dark-brown ones. At a population scale, darker birds had higher probability of colonization of the most isolated, empty sites. Explanations for species range expansion and population regulation usually make the inferred assumption that species traits are similar among individuals. However, in most species, there could be individual variation in niche requirements or dispersal propensities among individuals with different traits. Our results contribute to the growing appreciation that the individual traits, but not the average trait at the level of species, are important during population regulation and expansion processes.