Ecological traps and species distribution models: a challenge for prioritizing areas of conservation importance

Species distribution models analyse how species use different types of habitats. Their spatial predictions are often used to prioritize areas for conservation. Individuals may, however, prefer settling in habitat types of low quality compared to other available habitats. This ecological trap phenomenon is usually studied in a small number of habitat patches and consequences at the landscape level are largely unknown. It is therefore often unclear whether the spatial pattern of habitat use is aligned with the behavioural decisions made by the individuals during habitat selection or reflects actual variation in the quality of different habitat types. As species distribution models analyse the pattern of occurrence in different habitats, there is a conservation interest in examining what their predictions mean in terms of habitat quality when ecological traps are operating. Previous work in Belgium showed that red-backed shrikes Lanius collurio are more attracted to newly available clear-cut habitat in plantation forests than to the traditionally used farmland habitat. We developed models with shrike distribution data and compared their predictions with spatial variation in shrike reproductive performance used as a proxy for habitat quality. Models accurately predicted shrike distribution and identified the preferred clear-cut patches as the most frequently used habitat, but reproductive performance was lower in clear-cut areas than in farmland. With human-induced rapid environmental changes, organisms may indeed be attracted to low-quality habitats and occupy them at high densities. Consequently, the predictions of statistical models based on occurrence records may not align with variation in significant population parameters for the maintenance of the species. When species expand their range to novel habitats, such models are useful to document the spatial distribution of the organisms, but data on population growth rates are worth collecting before using model predictions to guide the spatial prioritization of conservation actions.     

Storing prey in larders affects nestling haematological condition in the Red-backed Shrike (Lanius collurio)

Many animals, including shrikes (Laniidae), are known to store food. This behaviour ensures food availability during inclement weather, for rearing nestlings or signalling territory quality. Unlike most other shrike species, which accumulate food before breeding, Red-backed Shrikes Lanius collurio store food in larders, mainly during the nestling and fledgling periods. If this is an adaptive behaviour, variation in this behaviour should affect either or both the number and condition of nestlings, which may benefit from such larders. We assessed the haematological condition of Red-backed Shrike nestlings in east-central Poland in 2021 using a suite of blood parameters relating to sites where shrikes did or did not store food. Haemoglobin concentrations were positively correlated with larders, being significantly higher in nestlings of pairs that stored prey compared with those whose parents did not do so. White blood cell counts were lower in nestlings whose parents kept larders. Red blood cell counts and glucose concentrations in nestlings were not related to whether parents had larders. Food storage probably ensures a more constant access to food, which results in a superior haematological condition in nestlings.     

Maladaptive habitat selection of a migratory passerine bird in a human-modified landscape

In human-altered environments, organisms may preferentially settle in poor-quality habitats where fitness returns are lower relative to available higher-quality habitats. Such ecological trapping is due to a mismatch between the cues used during habitat selection and the habitat quality. Maladaptive settlement decisions may occur when organisms are time-constrained and have to rapidly evaluate habitat quality based on incomplete knowledge of the resources and conditions that will be available later in the season. During a three-year study, we examined settlement decision-making in the long-distance migratory, open-habitat bird, the Red-backed shrike (Lanius collurio), as a response to recent land-use changes. In Northwest Europe, the shrikes typically breed in open areas under a management regime of extensive farming. In recent decades, Spruce forests have been increasingly managed with large-size cutblocks in even-aged plantations, thereby producing early-successional vegetation areas that are also colonised by the species. Farmland and open areas in forests create mosaics of two different types of habitats that are now occupied by the shrikes. We examined redundant measures of habitat preference (order of settlement after migration and distribution of dominant individuals) and several reproductive performance parameters in both habitat types to investigate whether habitat preference is in line with habitat quality. Territorial males exhibited a clear preference for the recently created open areas in forests with higher-quality males settling in this habitat type earlier. Reproductive performance was, however, higher in farmland, with higher nest success, offspring quantity, and quality compared to open areas in forests. The results showed strong among-year consistency and we can therefore exclude a transient situation. This study demonstrates a case of maladaptive habitat selection in a farmland bird expanding its breeding range to human-created open habitats in plantations. We discuss the reasons that could explain this decision-making and the possible consequences for the population dynamics and persistence.     

UV reflecting vole scent marks attract a passerine, the great grey shrike Lanius excubitor

Diurnal raptors have been shown to use ultraviolet vision and UV-reflecting vole scent marks to find suitable hunting areas. We studied if a passerine species, the great grey shrike Lanius excubitor that uses voles as its primary food, may also detect prey-patches in the same way. We conducted a laboratory experiment with ten shrikes. Each individual shrike had four options to choose from: (1) scent marks with UV light, (2) scent marks without UV light, (3) clean arena with UV light, and (4) clean arena without UV light. The birds preferred the scent-marked arena with UV light as measured by the number of scans and the time spent above it. Therefore, we suggest that great grey shrike probably uses UV cues to gain information on vole locations and abundance.     

Comparison of nest defence behaviour between two associate passerines

Nest predation is one of the most important factors limiting reproductive success, and antipredator behaviour can significantly reduce the loss of avian broods. I carried out field experiments on two sympatric passerines: the barred warbler and the red-backed shrike. Many authors have described the protective nature of nesting association between these species. However, we have little knowledge about the true nature of the relationships between associates. I examined (1) whether barred warblers and red-backed shrikes respond differently to an avian predator, and (2) whether males and females differ in the intensity of nest defence. Decoys of a known nest predator and a non-predatory control species were used to examine the types and relative intensity of parental response. I measured behavioural responsiveness by recording aggressive behaviour toward each model during the nestling period. Barred warblers and red-backed shrikes showed considerable variation in their response. Warblers more vigorously defended their own territories than shrikes. No differences between the sexes in antipredator behaviour in red-backed shrike were found. By contrast, in barred warbler, male was more involved in nest defence. The experimental tests provide evidence that these two species are able to differentiate between a predator and non-predator species.     

Using satellite imagery to assess breeding habitat availability of the endangered loggerhead shrike in Québec

The loggerhead shrike (Lanius ludovicianus) is a grassland bird species whose preferred nesting habitat in eastern Canada is pastureland. This species has been extirpated from much of its historical range in this region, and breeding habitat loss is suspected to be an important cause of this decline. We evaluated the availability of suitable breeding habitats in Québec using satellite imagery. Because this species no longer breeds in Québec, we established habitat selection criteria from known nesting sites in the adjacent province of Ontario, from analysis of a Landsat-TM satellite image, and applied these criteria to Landsat-TM images covering southern Québec. We developed regional landscape criteria in 100 km² plots and patch indices criteria at the pasture level. Spatial analyses were conducted to characterize plots and pastures on the basis of pasture availability and spatial distribution. Pastures suitable for nesting loggerhead shrikes were those fulfilling patch criteria at the pasture level and located in plots fulfilling regional landscape criteria. Overall, 310 out of 1700 plots located in the historical breeding range of the loggerhead shrike in Québec fulfilled landscape criteria, supporting 3988 pastures that fulfilled patch criteria. More than 500 of these pastures were visited to validate their current status. The Outaouais region would be the most suitable region for nesting loggerhead shrikes in southern Québec, where suitable breeding habitat still remains because more than two-thirds of visited sites were still pastureland, hawthorns were well-distributed in the region, and pastureland fragmentation was lowest. We conclude that the availability of breeding habitat does not limit the establishment of a breeding population of loggerhead shrike in southern Québec, as we estimated that thousands of hectares of suitable habitat still remain in that province.     

Species distribution models as a tool to estimate reproductive parameters: a case study with a passerine bird species

1.?Correlative species distribution models (SDMs) assess relationships between species distribution data and environmental features, to evaluate the environmental suitability (ES) of a given area for a species, by providing a measure of the probability of presence. If the output of SDMs represents the relationships between habitat features and species performance well, SDM results can be related also to other key parameters of populations, including reproductive parameters. To test this hypothesis, we evaluated whether SDM results can be used as a proxy of reproductive parameters (breeding output, territory size) in red-backed shrikes (Lanius collurio). 2.?The distribution of 726 shrike territories in Northern Italy was obtained through multiple focused surveys; for a subset of pairs, we also measured territory area and number of fledged juveniles. We used Maximum Entropy modelling to build a SDM on the basis of territory distribution. We used generalized least squares and spatial generalized mixed models to relate territory size and number of fledged juveniles to SDM suitability, while controlling for spatial autocorrelation. 3.?Species distribution models predicted shrike distribution very well. Territory size was negatively related to suitability estimated through SDM, while the number of fledglings significantly increased with the suitability of the territory. This was true also when SDM was built using only spatially and temporally independent data. 4.?Results show a clear relationship between ES estimated through presence-only SDMs and two key parameters related to species' reproduction, suggesting that suitability estimated by SDM, and habitat quality determining reproduction parameters in our model system, are correlated. Our study shows the potential use of SDMs to infer important fitness parameters; this information can have great importance in management and conservation.     

Density changes and habitat shift of great grey shrike <Emphasis Type="Italic">Lanius excubitor</Emphasis> during the non-breeding season

Bird population density and their behaviour during non-breeding periods are determined mainly by meteorological conditions which force birds to use alternative strategies — migration to areas with more advantageous conditions or adaptation to existing conditions. During the 2002/03–2004/5 non-breeding seasons in the open habitats of the Kotlina Orawsko-Nowotarska valley (S Poland) studies on population dynamics, behaviour and habitat use of great grey shrike were conducted. Results were grouped into four meteorological periods (autumn, early winter, winter and early spring). Density changes were seen between subsequent seasons (from 0.34 to 1.9 ind./10 km), as well as between meteorological periods (from 0.48 to 2.70 ind./10 km). Perching was the most frequently observed behaviour. The foraging habitats used changed in frequency. In subsequent meteorological periods a decrease of open habitats used was noted with a simultaneous increase in afforestations used. These changes did not depend on lasting snow cover. Changes in vegetation type used during hunting were not observed. Great grey shrikes most frequently used natural perches (trees, shrubs and high vegetation).     

Nest defense of the red-backed shrike Lanius collurio against five corvid species

We compared the antipredation behavior of the red-backed shrike against five European corvids including the jay, nutcracker, rook, crow, and raven. These species differ in body size and in the proportion of eggs and nestlings in their diets. The jay and nutcracker are the smallest, the rook and crow are middle-sized, and the raven being the largest corvid of all. The jay, crow, and raven are habitual nest predators, whereas the nutcracker and rook are not. The harmless pigeon was presented as a control. We analyzed (1) the number of attacks executed by shrikes against intruder presented close to shrike nests and (2) the distance between the intruder and the shrikes during the trial. The small corvids (the jay and nutcracker) were attacked significantly more intensively than the other, larger, corvids (the rook, crow, and raven) and pigeon control. All three large corvids were attacked as exceptionally as the pigeon. Shrikes approached closer to the small corvids and the pigeon than to the large corvids. These results imply that shrike antipredation strategy is adjusted to intruder size, but not to the level of intruder nest plundering specialization. Shrikes weigh up their ability to chase a given intruder away and avoid pointless aggression against large, undefeatable, intruders. This suggests that shrikes are able to asses not only the dangerousness of the intruder but also the potential advantageousness, or otherwise, of active defense.     

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.     

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.     

Space Use in Response to Conspecifics by the Red-backed Salamander (Plethodon cinereus, Plethodontidae, Caudata)

The presence of conspecifics, as well as the distribution and quality of resources, can influence habitat settlement decisions of animals. We studied the habitat preference and group size of the red-backed salamander, Plethodon cinereus , from the Upper Peninsula of Michigan. In contrast to previous work with P. cinereus from Virginia, we found groups of P. cinereus within higher quality habitat than solitary individuals and these groups contained both small and large individuals. Under identical conditions in the laboratory, salamanders from Virginia formed a uniform spatial distribution and individuals from northern Michigan displayed an aggregated spatial distribution. Our results suggest that P. cinereus from northern Michigan preferentially settle near conspecifics, while P. cinereus from Virginia avoid them. Our laboratory experiments also suggest that the aggregation behaviour of P. cinereus from northern Michigan may be the result of attraction to conspecifics rather than attraction to habitat features.     

Population structure of loggerhead shrikes in the California Channel Islands

The loggerhead shrike (Lanius ludovicianus), a songbird that hunts like a small raptor, maintains breeding populations on seven of the eight California Channel Islands. One of the two subspecies, L. l. anthonyi, was described as having breeding populations on six of the islands while a second subspecies, L. l. mearnsi, was described as being endemic to San Clemente Island. Previous genetic studies have demonstrated that the San Clemente Island loggerhead shrike is well differentiated genetically from both L. l. anthonyi and mainland populations, despite the fact that birds from outside the population are regular visitors to the island. Those studies, however, did not include a comparison between San Clemente Island shrikes and the breeding population on Santa Catalina Island, the closest island to San Clemente. Here we use mitochondrial control region sequences and nuclear microsatellites to investigate the population structure of loggerhead shrikes in the Channel Islands. We confirm the genetic distinctiveness of the San Clemente Island loggerhead shrike and, using Bayesian clustering analysis, demonstrate the presence and infer the source of the nonbreeding visitors. Our results indicate that Channel Island loggerhead shrikes comprise three distinct genetic clusters that inhabit: (i) San Clemente Island, (ii) Santa Catalina Island and (iii) the Northern Channel Islands and nearby mainland; they do not support a recent suggestion that all Channel Island loggerhead shrikes should be managed as a single entity.     

Are the birds dangerous for insect pollinators? The relationship between hymenopterans and the red-backed shrike

Studies of the relationships between insectivorous birds and insects have focused on woodland habitats, where insects have usually been regarded as tree pests. But some insects are of the utmost importance to the human economy, especially hymenopterans, which pollinate plants such as fruit trees in orchards. Here we report on a 5-year study of the influence of the red-backed shrike Lanius collurio, a territorial insectivorous bird, on the numbers of hymenopterans in the extensively farmed landscape of eastern Poland. The relevant relationships were based on an analysis of the shrike’s diet and the numbers of hymenopterans in and beyond nesting locations. There was a weak but statistically significant relationship between the proportion of hymenopterans in the red-backed shrike’s diet and the place and time of prey capture, but this was dependent solely on the location of the shrike’s territory. No relationship was found between the numbers of hymenopterans and the presence of the red-backed shrike, agricultural site type or season effect. In conclusion, territorial bird species that occur in low densities probably have only a marginal influence on hymenopterans, including pollinators.     

Traditional versus non-traditional nest-site choice: alternative decision strategies for nest-site selection

In order to understand habitat selection, it is important to consider the way individual animals assess the suitability of a future reproductive site. One way of investigating mechanisms (such as those involved in nest site selection) is to examine breeding success and habitat characteristics in terms of animals returning to a place where they have already reproduced and using the same location over successive years or searching for new alternatives. This approach seems especially suitable for testing recent hypotheses suggesting that nest site selection is an integrative process that includes the use of social information (e.g. past breeding success of conspecifics). Determining the factors that elicit conservative or innovative behaviour regarding nest-site selection could be important for improving our understanding of habitat selection decisions in animals. More than half of the nests of the long-distance migratory lesser grey shrike Lanius minor, are built in the same or neighbouring trees. We found no evidence that habitat characteristics influence nest-site tradition. On the contrary, social information in terms of the presence of conspecifics and past reproductive success in terms of complete nest failures due to nest predation (but not detailed information such as variation in fledgling number) influenced nest-site tradition. Hence, social information and past reproductive success may play a role in nest-site choice in this species. Our results further demonstrate that previous experience with a nest site does not appear to be beneficial.     

Who is in the neighborhood? Conspecific and heterospecific responses to perceived density for breeding habitat selection

Theoretical models of habitat selection often incorporate negative density dependence. Despite strong negative density‐dependent effects on habitat selection, more recent studies indicate that animals settle near members of their own (conspecific) and other species (heterospecific) when selecting habitat with social cues. Social cue use for habitat selection is particularly common among songbirds, but few studies have investigated if songbirds use social cues to assess conspecific or heterospecific density (as opposed to just presence/absence) when making settlement decisions. We conducted a playback experiment to evaluate if yellow warblers (Setophaga petechia) and willow flycatchers (Empidonax traillii), two potential competitors for breeding habitat, use social cues to assess density (conspecific for warblers and heterospecific for flycatchers) when selecting breeding locations at two spatial scales. We simulated yellow warbler density to be high or low at multiple treatment plots (3.14 ha) with song playback and then evaluated settlement decisions by comparing yellow warbler and willow flycatcher abundances across plots (broad‐scale habitat selection) and individual space use within plots (fine‐scale territory establishment). Yellow warbler density treatments did not affect habitat selection by yellow warblers at the broad scale, but caused individuals to cluster territories at high‐density treatments. Willow flycatchers were most abundant at high‐density treatment plots, but yellow warbler density treatments did not affect territory locations. The results indicate that perceived density affects the habitat selection process for both conspecifics and heterospecifics.     

Feather deuterium measurements reveal origins of migratory western loggerhead shrikes (Lanius ludovicianus excubitorides) wintering in Mexico

Understanding the winter distributions of migrant birds is important because productivity and recruitment are influenced by conditions at several locations and periods in the life cycle of individuals. The western loggerhead shrike, Lanius ludovicianus excubitorides , is a threatened species in Canada, and its decline is attributed to potential limitations on the wintering grounds. We examined patterns of stable-hydrogen isotope (δD) distributions in feathers of loggerhead shrikes, primarily of L. l. excubitorides , during winter at three regions in north and central Mexico, to establish relative abundance and origins of migrants. We also investigated potential movements of Mexican winter resident individuals. Using shrike museum specimens of known summer provenance, a shrike deuterium base map for Mexico was developed from isotopic measurement of feathers of resident shrikes and use of a recently established feather base map for raptors in North America. Stable hydrogen isotope analyses of inner secondary feather (s9) of all loggerhead shrikes examined in Mexico during winter indicated that north-central (Region A), north-eastern (Region B) and south-central (Region C) sites in Mexico consisted of 28.1%, 73.7% and 63.8% of migrant individuals from northern breeding grounds, respectively. Isotopic evidence suggested movements of a few local residents birds (7.9%) into the Chihuahuan desert from south-western USA and north-eastern Mexico to winter.     

Ecological and social effects on reproduction and local recruitment in the red-backed shrike

Numerous hypotheses have been proposed to explain variation in reproductive performance and local recruitment of animals. While most studies have examined the influence of one or a few social and ecological factors on fitness traits, comprehensive analyses jointly testing the relative importance of each of many factors are rare. We investigated how a multitude of environmental and social conditions simultaneously affected reproductive performance and local recruitment of the red-backed shrike Lanius collurio (L.). Specifically, we tested hypotheses relating to timing of breeding, parental quality, nest predation, nest site selection, territory quality, intraspecific density and weather. Using model selection procedures, predictions of each hypothesis were first analysed separately, before a full model was constructed including variables selected in the single-hypothesis tests. From 1988 to 1992, 50% of 332 first clutches produced at least one fledgling, while 38.7% of 111 replacement clutches were successful. Timing of breeding, nest site selection, predation pressure, territory quality and intraspecific density influenced nest success in the single-hypothesis tests. The full model revealed that nest success was negatively associated with laying date, intraspecific density, and year, while nest success increased with nest concealment. Number of fledglings per successful nest was only influenced by nest concealment: better-camouflaged nests produced more fledglings. Probability of local recruitment was related to timing of breeding, parental quality and territory quality in the single-hypothesis tests. The full models confirmed the important role of territory quality for recruitment probability. Our results suggest that reproductive performance, and particularly nest success, of the red-backed shrike is primarily affected by timing of breeding, nest site selection, and intraspecific density. This study highlights the importance of considering many factors at the same time, when trying to evaluate their relative contributions to fitness and life history evolution.Electronic supplementary material Supplementary material is available for this article at     

Social environment affects juvenile dispersal in great tits (Parus major)

1.?Habitat selection can affect individual fitness, and therefore, individuals are expected to assess habitat quality of potential breeding sites before settlement. 2.?We investigated the role of social environment on juvenile dispersal behaviour in the great tit (Parus major). Two main contradictory hypotheses can be formulated regarding social effects on juvenile dispersal as follows: (i) High fledgling density and sex ratio may enhance the intensity of local (kin) competition and, therefore, reduce individual survival chance, enhance emigration and reduce settlement ('repulsion' hypothesis) (ii) Alternatively, high fledgling density and sex ratio may signal high-quality habitat or lead to aggregation and thus increase individual survival chance, reduce emigration and enhance settlement ('attraction' hypothesis). 3.?To disentangle positive from negative effects of high density and male-biased sex ratio on dispersal, we manipulated the social composition of the fledgling population in 12 semi-isolated nest-box areas (plots) via a change of fledgling density (low/high) as well as fledgling sex ratio (female-biased/balanced/male-biased) across 3?years. We then tested whether experimental variation in male and female fledgling densities affected variation in local survival, emigration and settlement of juveniles, and whether social effects on survival and dispersal support the 'repulsion' or 'attraction' hypothesis. 4.?We found no experimental effects on local survival and emigration probabilities. However, consistent with the 'attraction' hypothesis, settlement was significantly and positively affected by local experimental sex ratio in each of the study years: both male and female juveniles avoided female-biased plots and settled more in plots that were balanced and male-biased the previous year. 5.?Our study provides unprecedented experimental evidence that local sex ratio plays a causal role in habitat selection. We suggest that settlers avoid female-biased plots because a high proportion of females may reflect the absence or the low quality of local resources in the habitat. Alternatively, male territory acquisition may be facilitated by a high local density of 'candidate' males, and therefore, juveniles were less successful in settling in female-biased plots.     

Modelling spatial and environmental effects at the edge of the distribution: the red-backed shrike Lanius collurio in Northern Portugal

This study examined the interplay of spatial and environmental effects shaping the range margin of the red‐backed shrike (Lanius collurio) in northern Portugal. The occurrence of shrikes in 10 × 10 km UTM squares was related to three sets of explanatory variables, reflecting environmental effects (climate and habitat), large‐scale spatial trends, and neighbourhood influences (considering an autologistic term); spatial variables were used as surrogates for historical and demographic factors. Multiple logistic regression models were built for each set, and then variation partitioning based on partial regressions isolated the unique and shared components of explained variation. The environmental model revealed a dominant influence of climate effects, with the occurrence of shrikes increasing with frost and thermal amplitude, declining with insolation, and responding unimodally to rainfall. There was a weaker influence of habitat conditions, though shrikes were more likely with increasing cover by annual crops and pastures, and decreasing forest cover. Only a relatively small proportion of explained variation was due to a ‘pure’ environmental component (10.4%), as most variation explained by environmental factors appeared spatially structured (51.9%). The unique contributions of spatial variables to the overall model were also small, though the neighbourhood effects appeared relatively stronger than large‐scale trends. Taken together, results suggested that the south‐western range margin of the red‐backed shrike was largely determined by spatially structured environmental factors. Nevertheless, there were also ‘pure’ environmental factors determining some isolate occurrences irrespective of any spatial structure, and ‘pure’ spatial factors that appeared to favour the occupation of squares surrounding the core distribution areas irrespective of environmental conditions. These results add to the growing evidence that both environmental and spatial factors need to be considered in predictive modelling of species range margins.