Unclear relationships between mean survival rate and its environmental variance in vertebrates.

Current environmental changes may increase temporal variability of life history traits of species thus affecting their long-term population growth rate and extinction risk. If there is a general relationship between environmental variances (EVs) and mean annual survival rates of species, that relationship could be used as a guideline for analyses of population growth and extinction risk for populations, where data on EVs are missing. For this purpose, we present a comprehensive compilation of 252 EV estimates from 89 species belonging to five vertebrate taxa (birds, mammals, reptiles, amphibians and fish) covering mean annual survival rates from 0.01 to 0.98. Since variances of survival rates are constrained by their means, particularly for low and high mean survival rates, we assessed whether any observed relationship persisted after applying two types of commonly used variance stabilizing transformations: relativized EVs (observed/mathematical maximum) and logit-scaled EVs. With raw EVs at the arithmetic scale, mean-variance relationships of annual survival rates were hump-shaped with small EVs at low and high mean survival rates and higher (and widely variable) EVs at intermediate mean survival rates. When mean annual survival rates were related to relativized EVs the hump-shaped pattern was less distinct than for raw EVs. When transforming EVs to logit scale the relationship between mean annual survival rates and EVs largely disappeared. The within-species juvenile-adult slopes were mainly positive at low (<0.5) and negative at high (>0.5) mean survival rates for raw and relativized variances while these patterns disappeared when EVs were logit transformed. Uncertainties in how to interpret the results of relativized and logit-scaled EVs, and the observed high variation in EV's for similar mean annual survival rates illustrates that extrapolations of observed EVs and tests of life history drivers of survival-EV relationships need to also acknowledge the large variation in these parameters.

Partitioning variance in population growth for models with environmental and demographic stochasticity.

How demographic factors lead to variation or change in growth rates can be investigated using life table response experiments (LTRE) based on structured population models. Traditionally, LTREs focused on decomposing the asymptotic growth rate, but more recently decompositions of annual 'realized' growth rates using 'transient' LTREs have gained in popularity. Transient LTREs have been used particularly to understand how variation in vital rates translate into variation in growth for populations under long-term study. For these, complete population models may be constructed to investigate how temporal variation in environmental drivers affect vital rates. Such investigations have usually come down to estimating covariate coefficients for the effects of environmental variables on vital rates, but formal ways of assessing how they lead to variation in growth rates have been lacking. We extend transient LTREs to further partition the contributions from vital rates into contributions from temporally varying factors that affect them. The decomposition allows one to compare the resultant effect on the growth rate of different environmental factors, as well as density dependence, which may each act via multiple vital rates. We also show how realized growth rates can be decomposed into separate components from environmental and demographic stochasticity. The latter is typically omitted in LTRE analyses. We illustrate these extensions with an integrated population model (IPM) for data from a 26 years study on northern wheatears (Oenanthe oenanthe), a migratory passerine bird breeding in an agricultural landscape. For this population, consisting of around 50-120 breeding pairs per year, we partition variation in realized growth rates into environmental contributions from temperature, rainfall, population density and unexplained random variation via multiple vital rates, and from demographic stochasticity. The case study suggests that variation in first year survival via the unexplained random component, and adult survival via temperature are two main factors behind environmental variation in growth rates. More than half of the variation in growth rates is suggested to come from demographic stochasticity, demonstrating the importance of this factor for populations of moderate size.

The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Temporal trends in opportunistic citizen science reports across multiple taxa.

Opportunistic reporting of species observations to online platforms provide one of the most extensive sources of information about the distribution and status of organisms in the wild. The lack of a clear sampling design, and changes in reporting over time, leads to challenges when analysing these data for temporal change in organisms. To better understand temporal changes in reporting, we use records submitted to an online platform in Sweden (Artportalen), currently containing 80 million records. Focussing on five taxonomic groups, fungi, plants, beetles, butterflies and birds, we decompose change in reporting into long-term and seasonal trends, and effects of weekdays, holidays and weather variables. The large surge in number of records since the launch of the, initially taxa-specific, portals is accompanied by non-trivial long-term and seasonal changes that differ between the taxonomic groups and are likely due to changes in, and differences between, the user communities and observer behaviour.

Fluctuating optimum and temporally variable selection on breeding date in birds and mammals.

Temporal variation in natural selection is predicted to strongly impact the evolution and demography of natural populations, with consequences for the rate of adaptation, evolution of plasticity, and extinction risk. Most of the theory underlying these predictions assumes a moving optimum phenotype, with predictions expressed in terms of the temporal variance and autocorrelation of this optimum. However, empirical studies seldom estimate patterns of fluctuations of an optimum phenotype, precluding further progress in connecting theory with observations. To bridge this gap, we assess the evidence for temporal variation in selection on breeding date by modeling a fitness function with a fluctuating optimum, across 39 populations of 21 wild animals, one of the largest compilations of long-term datasets with individual measurements of trait and fitness components. We find compelling evidence for fluctuations in the fitness function, causing temporal variation in the magnitude, but not the direction of selection. However, fluctuations of the optimum phenotype need not directly translate into variation in selection gradients, because their impact can be buffered by partial tracking of the optimum by the mean phenotype. Analyzing individuals that reproduce in consecutive years, we find that plastic changes track movements of the optimum phenotype across years, especially in bird species, reducing temporal variation in directional selection. This suggests that phenological plasticity has evolved to cope with fluctuations in the optimum, despite their currently modest contribution to variation in selection.

Cannot see the diversity for all the species: Evaluating inclusion criteria for local species lists when using abundant citizen science data.

Abundant citizen science data on species occurrences are becoming increasingly available and enable identifying composition of communities occurring at multiple sites with high temporal resolution. However, for species displaying temporary patterns of local occurrences that are transient to some sites, biodiversity measures are clearly dependent on the criteria used to include species into local species lists. Using abundant opportunistic citizen science data from frequently visited wetlands, we investigated the sensitivity of α- and ß-diversity estimates to the use raw versus detection-corrected data and to the use of inclusion criteria for species presence reflecting alternative site use. We tested seven inclusion criteria (with varying number of days required to be present) on time series of daily occurrence status during a breeding season of 90 days for 77 wetland bird species. We show that even when opportunistic presence-only observation data are abundant, raw data may not produce reliable local species richness estimates and rank sites very differently in terms of species richness. Furthermore, occupancy model based α- and ß-diversity estimates were sensitive to the inclusion criteria used. Total species lists (all species observed at least once during a season) may therefore mask diversity differences among sites in local communities of species, by including vagrant species on potentially breeding communities and change the relative rank order of sites in terms of species richness. Very high sampling effort does not necessarily free opportunistic data from its inherent bias and can produce a pattern in which many species are observed at least once almost everywhere, thus leading to a possible paradox: The large amount of biological information may hinder its usefulness. Therefore, when prioritizing among sites to manage or preserve species diversity estimates need to be carefully related to relevant inclusion criteria depending on the diversity estimate in focus.

Why we should care about movements: Using spatially explicit integrated population models to assess habitat source-sink dynamics.

Assessing the source-sink status of populations and habitats is of major importance for understanding population dynamics and for the management of natural populations. Sources produce a net surplus of individuals (per capita contribution to the metapopulation > 1) and will be the main contributors for self-sustaining populations, whereas sinks produce a deficit (contribution < 1). However, making these types of assessments is generally hindered by the problem of separating mortality from permanent emigration, especially when survival probabilities as well as moved distances are habitat-specific. To address this long-standing issue, we propose a spatial multi-event integrated population model (IPM) that incorporates habitat-specific dispersal distances of individuals. Using information about local movements, this IPM adjusts survival estimates for emigration outside the study area. Analysing 24 years of data on a farmland passerine (the northern wheatear Oenanthe oenanthe), we assessed habitat-specific contributions, and hence the source-sink status and temporal variation of two key breeding habitats, while accounting for habitat- and sex-specific local dispersal distances of juveniles and adults. We then examined the sensitivity of the source-sink analysis by comparing results with and without accounting for these local movements. Estimates of first-year survival, and consequently habitat-specific contributions, were higher when local movement data were included. The consequences from including movement data were sex specific, with contribution shifting from sink to likely source in one habitat for males, and previously noted habitat differences for females disappearing. Assessing the source-sink status of habitats is extremely challenging. We show that our spatial IPM accounting for local movements can reduce biases in estimates of the contribution by different habitats, and thus reduce the overestimation of the occurrence of sink habitats. This approach allows combining all available data on demographic rates and movements, which will allow better assessment of source-sink dynamics and better informed conservation interventions.

Can Mitogenomes of the Northern Wheatear (Oenanthe oenanthe) Reconstruct Its Phylogeography and Reveal the Origin of Migrant Birds?

The Northern Wheatear (Oenanthe oenanthe, including the nominate and the two subspecies O. o. leucorhoa and O. o. libanotica) and the Seebohm's Wheatear (Oenanthe seebohmi) are today regarded as two distinct species. Before, all four taxa were regarded as four subspecies of the Northern Wheatear. Their classification has exclusively been based on ecological and morphological traits, while their molecular characterization is still missing. With this study, we used next-generation sequencing to assemble 117 complete mitochondrial genomes covering O. o. oenanthe, O. o. leucorhoa and O. seebohmi. We compared the resolution power of each individual mitochondrial marker and concatenated marker sets to reconstruct the phylogeny and estimate speciation times of three taxa. Moreover, we tried to identify the origin of migratory wheatears caught on Helgoland (Germany) and on Crete (Greece). Mitogenome analysis revealed two different ancient lineages that separated around 400,000 years ago. Both lineages consisted of a mix of subspecies and species. The phylogenetic trees, as well as haplotype networks are incongruent with the present morphology-based classification. Mitogenome could not distinguish these presumed species. The genetic panmixia among present populations and taxa might be the consequence of mitochondrial introgression between ancient wheatear populations.

Weak effects of geolocators on small birds: A meta-analysis controlled for phylogeny and publication bias.

Currently, the deployment of tracking devices is one of the most frequently used approaches to study movement ecology of birds. Recent miniaturization of light-level geolocators enabled studying small bird species whose migratory patterns were widely unknown. However, geolocators may reduce vital rates in tagged birds and may bias obtained movement data. There is a need for a thorough assessment of the potential tag effects on small birds, as previous meta-analyses did not evaluate unpublished data and impact of multiple life-history traits, focused mainly on large species and the number of published studies tagging small birds has increased substantially. We quantitatively reviewed 549 records extracted from 74 published and 48 unpublished studies on over 7,800 tagged and 17,800 control individuals to examine the effects of geolocator tagging on small bird species (body mass <100 g). We calculated the effect of tagging on apparent survival, condition, phenology and breeding performance and identified the most important predictors of the magnitude of effect sizes. Even though the effects were not statistically significant in phylogenetically controlled models, we found a weak negative impact of geolocators on apparent survival. The negative effect on apparent survival was stronger with increasing relative load of the device and with geolocators attached using elastic harnesses. Moreover, tagging effects were stronger in smaller species. In conclusion, we found a weak effect on apparent survival of tagged birds and managed to pinpoint key aspects and drivers of tagging effects. We provide recommendations for establishing matched control group for proper effect size assessment in future studies and outline various aspects of tagging that need further investigation. Finally, our results encourage further use of geolocators on small bird species but the ethical aspects and scientific benefits should always be considered.

Factors influencing plasticity in the arrival-breeding interval in a migratory species reacting to climate change.

Climate change is profoundly affecting the phenology of many species. In migratory birds, there is evidence for advances in their arrival time at the breeding ground and their timing of breeding, yet empirical studies examining the interdependence between arrival and breeding time are lacking. Hence, evidence is scarce regarding how breeding time may be adjusted via the arrival-breeding interval to help local populations adapt to local conditions or climate change. We used long-term data from an intensively monitored population of the northern wheatear (Oenanthe oenanthe) to examine the factors related to the length of 734 separate arrival-to-breeding events from 549 individual females. From 1993 to 2017, the mean arrival and egg-laying dates advanced by approximately the same amount (~5-6 days), with considerable between-individual variation in the arrival-breeding interval. The arrival-breeding interval was shorter for: (a) individuals that arrived later in the season compared to early-arriving birds, (b) for experienced females compared to first-year breeders, (c) as spring progressed, and (d) in later years compared to earlier ones. The influence of these factors was much larger for birds arriving earlier in the season compared to later arriving birds, with most effects on variation in the arrival-breeding interval being absent in late-arriving birds. Thus, in this population it appears that the timing of breeding is not constrained by arrival for early- to midarriving birds, but instead is dependent on local conditions after arrival. For late-arriving birds, however, the timing of breeding appears to be influenced by arrival constraints. Hence, impacts of climate change on arrival dates and local conditions are expected to vary for different parts of the population, with potential negative impacts associated with these factors likely to differ for early- versus late-arriving birds.

Quantifying the links between land use and population growth rate in a declining farmland bird.

Land use is likely to be a key driver of population dynamics of species inhabiting anthropogenic landscapes, such as farmlands. Understanding the relationships between land use and variation in population growth rates is therefore critical for the management of many farmland species. Using 24 years of data of a declining farmland bird in an integrated population model, we examined how spatiotemporal variation in land use (defined as habitats with "Short" and "Tall" ground vegetation during the breeding season) and habitat-specific demographic parameters relates to variation in population growth taking into account individual movements between habitats. We also evaluated contributions to population growth using transient life table response experiments which gives information on contribution of past variation of parameters and real-time elasticities which suggests future scenarios to change growth rates. LTRE analyses revealed a clear contribution of Short habitats to the annual variation in population growth rate that was mostly due to fledgling recruitment, whereas there was no evidence for a contribution of Tall habitats. Only 18% of the variation in population growth was explained by the modeled local demography, the remaining variation being explained by apparent immigration (i.e., the residual variation). We discuss potential biological and methodological reasons for high contributions of apparent immigration in open populations. In line with LTRE analysis, real-time elasticity analysis revealed that demographic parameters linked to Short habitats had a stronger potential to influence population growth rate than those of Tall habitats. Most particularly, an increase of the proportion of Short sites occupied by Old breeders could have a distinct positive impact on population growth. High-quality Short habitats such as grazed pastures have been declining in southern Sweden. Converting low-quality to high-quality habitats could therefore change the present negative population trend of this, and other species with similar habitat requirements.

Hatching failure and accumulation of organic pollutants through the terrestrial food web of a declining songbird in Western Europe.

Population growth in passerine birds is largely driven by fecundity. If fecundity is affected, for instance by hatching failure, populations may decline. We noted high hatching failure of up to 27% per year in relict populations of the Northern wheatear (Oenanthe oenanthe) in The Netherlands, a strongly declining, migratory passerine in Europe. This hatching failure itself can cause population decline, irrespective of other adverse factors. Additionally, we investigated the cause of hatching failure. Unhatched eggs showed egg yolk infections or embryonic malformations, part of which is associated with the actions of dioxin-like compounds (DLCs). Indeed, DLCs appear to bioaccumulate in the local foodweb, where the soil contained only background concentrations, similar to those found at many other locations. DLC concentrations in Dutch eggs were six-fold higher than those in a reference population in Sweden, where egg failure was only 6%. However, Northern wheatears appear to be only moderately sensitive to the actions of DLCs, because of their specific Ah-receptor type which may moderate the receptor mediated effects of DLCs. This indicates that the concentrations of DLCs, although elevated, may not have caused the embryo malformations or the low hatching rates. We discuss whether other toxins may be important or imbalances in the nutrition and if inbreeding may play a larger role than expected.

Disentangling the effects of date, individual, and territory quality on the seasonal decline in fitness.

The seasonal timing of reproduction is a major fitness factor in many organisms. Commonly, individual fitness declines with time in the breeding season. We investigated three suggested but rarely tested hypotheses for this seasonal fitness decline: (1) time per se (date hypothesis), (2) late breeders are of lower quality than early ones (individual quality hypothesis), and (3) late breeders are breeding at poorer territories than early breeders (territory quality hypothesis). We used Bayesian variance component analyses to examine reproductive output (breeding success, number fledged, and number of recruits) from repeated observations of female Northern Wheatears (Oenanthe oenanthe) and individual territories from a 20-yr population study. The major part of the observed seasonal decline in reproductive output seemed to be driven by date-related effects, whereas female age and territory type (i.e., known indicators of temporary quality) contributed to a smaller degree. Other, persistent effects linked to individual and territory identity did not show any clear patterns on the seasonal decline in reproductive output. To better disentangle the quality effects (persistent and temporary) of individual and territory from effects caused by the deterioration of the environment we suggest a protocol combining experimental manipulation of breeding time with a variance-covariance partitioning method used here.

Marked reduction in demographic rates and reduced fitness advantage for early breeding is not linked to reduced thermal matching of breeding time.

Warmer springs may cause animals to become mistimed if advances of spring timing, including available resources and of timing of breeding occur at different speed. We used thermal sums (cumulative sum of degree days) during spring to describe the thermal progression (timing) of spring and investigate its relationship to breeding phenology and demography of a long-distant migrant bird, the northern wheatear (Oenanthe oenanthe L.). We first compare 20-year trends in spring timing, breeding time, selection for breeding time, and annual demographic rates. We then explicitly test whether annual variation in selection for breeding time and demographic rates associates with the degree of phenological matching between breeding time and thermal progression of spring. Both thermal progression of spring and breeding time of wheatears advanced in time during the study period. But despite breeding on average 7 days earlier with respect to date, wheatears bred about 4 days later with respect to thermal spring progression. Over the same time period, selection for breeding time changed from distinct within-season advantage of breeding early to no or very weak advantage. Furthermore, demographic rates (nest success, fledgling production, recruitment, adult survival) and nestling weight declined markedly by 16%-79%. Those temporal trends suggest that a reduced degree of phenological matching may affect within-season fitness advantage of early breeding and population demographic rates. In contrast, when we investigate links based on annual variation, we find no significant relationship between either demographic rates or fitness advantage of early breeding with annual variation in the degree of phenological matching. Our results show that corresponding temporal trends in phenological matching, selection for breeding time and demographic rates are inconclusive evidence for demographic effects of changed phenological matching. Instead, we suggest that the trends in selection for breeding time and demographic rates are due to a general deterioration of the breeding environment.

Rainfall during parental care reduces reproductive and survival components of fitness in a passerine bird.

Adverse weather conditions during parental care may have direct consequences for offspring production, but longer-term effects on juvenile and parental survival are less well known. We used long-term data on reproductive output, recruitment, and parental survival in northern wheatears (Oenanthe oenanthe) to investigate the effects of rainfall during parental care on fledging success, recruitment success (juvenile survival), and parental survival, and how these effects related to nestling age, breeding time, habitat quality, and parental nest visitation rates. While accounting for effects of temperature, fledging success was negatively related to rainfall (days > 10 mm) in the second half of the nestling period, with the magnitude of this effect being greater for breeding attempts early in the season. Recruitment success was, however, more sensitive to the number of rain days in the first half of the nestling period. Rainfall effects on parental survival differed between the sexes; males were more sensitive to rain during the nestling period than females. We demonstrate a probable mechanism driving the rainfall effects on reproductive output: Parental nest visitation rates decline with increasing amounts of daily rainfall, with this effect becoming stronger after consecutive rain days. Our study shows that rain during the nestling stage not only relates to fledging success but also has longer-term effects on recruitment and subsequent parental survival. Thus, if we want to understand or predict population responses to future climate change, we need to consider the potential impacts of changing rainfall patterns in addition to temperature, and how these will affect target species' vital rates.

Decomposing the seasonal fitness decline.

Seasonal fitness declines are common, but the relative contribution of different reproductive components to the seasonal change in the production of reproductive young, and the component-specific drivers of this change is generally poorly known. We used long-term data (17 years) on breeding time (i.e. date of first egg laid) in northern wheatears (Oenanthe oenanthe) to investigate seasonal reproductive patterns and estimate the relative contributions of reproductive components to the overall decline in reproduction, while accounting for factors potentially linked to seasonal declines, i.e. individual and habitat quality. All reproductive components-nest success (reflecting nest predation rate), clutch size, fledging success and recruitment success-showed a clear decline with breeding time whereas subsequent adult survival did not. A non-linear increase in nest predation rate caused nest success to decline rapidly early in the season and level off at ~80% success late in the breeding season. The combined seasonal decline in all reproductive components caused the mean production of recruits per nest to drop from around 0.7-0.2; with the relative contribution greatest for recruitment success which accounted for ~50% of the decline. Our data suggest that changing environmental conditions together with effects of nest predation have strong effects on the seasonal decline in fitness. Our demonstration of the combined effects of all reproductive components and their relative contribution shows that omitting data from later stages of breeding (recruitment) can greatly underestimate seasonal fitness declines.

Effect of geolocators on migration and subsequent breeding performance of a long-distance passerine migrant.

Geolocators are small light-weight data loggers used to track individual migratory routes, and their use has increased exponentially in birds. However, the effects of geolocators on individual performance are still poorly known. We studied geolocator effects on a long-distance migrating passerine bird, the northern wheatear (Oenanthe oenanthe L.). We asked the general question of whether geolocators affect migratory behaviour and subsequent reproductive performance of small passerines by comparing arrival time, breeding time, breeding success and survival of geolocator versus control birds of known identity and breeding history. During two years geolocator birds (n=37) displayed a lower apparent survival (30%) as compared to controls (45%, n=164). Furthermore, returning geolocator birds (n=12) arrived on average 3.5 days later, started laying eggs 6.3 days later, and had lower nest success (25%) than control birds (78%). Our results suggest that geolocators affect migratory performance with carry-over effects to the timing of breeding and reproductive success in the subsequent breeding season. We discuss the implications of such geolocator effects for the study of migratory strategies of small passerines in general and suggest how to identify and investigate such effects in the future.

The short- and long-term fitness consequences of natal dispersal in a wild bird population.

Dispersal is a key process in population and evolutionary ecology. Individual decisions are affected by fitness consequences of dispersal, but these are difficult to measure in wild populations. A long-term dataset on a geographically closed bird population, the Mauritius kestrel, offers a rare opportunity to explore fitness consequences. Females dispersed further when the availability of local breeding sites was limited, whereas male dispersal correlated with phenotypic traits. Female but not male fitness was lower when they dispersed longer distances compared to settling close to home. These results suggest a cost of dispersal in females. We found evidence of both short- and long-term fitness consequences of natal dispersal in females, including reduced fecundity in early life and more rapid aging in later life. Taken together, our results indicate that dispersal in early life might shape life history strategies in wild populations.

Contrast in edge vegetation structure modifies the predation risk of natural ground nests in an agricultural landscape.

Nest predation risk generally increases nearer forest-field edges in agricultural landscapes. However, few studies test whether differences in edge contrast (i.e. hard versus soft edges based on vegetation structure and height) affect edge-related predation patterns and if such patterns are related to changes in nest conspicuousness between incubation and nestling feeding. Using data on 923 nesting attempts we analyse factors influencing nest predation risk at different edge types in an agricultural landscape of a ground-cavity breeding bird species, the Northern Wheatear (Oenanthe oenanthe). As for many other bird species, nest predation is a major determinant of reproductive success in this migratory passerine. Nest predation risk was higher closer to woodland and crop field edges, but only when these were hard edges in terms of ground vegetation structure (clear contrast between tall vs short ground vegetation). No such edge effect was observed at soft edges where adjacent habitats had tall ground vegetation (crop, ungrazed grassland). This edge effect on nest predation risk was evident during the incubation stage but not the nestling feeding stage. Since wheatear nests are depredated by ground-living animals our results demonstrate: (i) that edge effects depend on edge contrast, (ii) that edge-related nest predation patterns vary across the breeding period probably resulting from changes in parental activity at the nest between the incubation and nestling feeding stage. Edge effects should be put in the context of the nest predator community as illustrated by the elevated nest predation risk at hard but not soft habitat edges when an edge is defined in terms of ground vegetation. These results thus can potentially explain previously observed variations in edge-related nest predation risk.

Prospectors combine social and environmental information to improve habitat selection and breeding success in the subsequent year.

1. Because habitats have profound effects on individual fitness, there is strong selection for improving the choice of breeding habitat. One possible mechanism is for individuals to use public information when prospecting future breeding sites; however, to our knowledge, no study has shown prospecting behaviour to be directly linked to subsequent choice of breeding site and future reproductive success. 2. We collected long-term data on territory-specific prospecting behaviour and subsequent breeding in the short-lived northern wheatear (Oenanthe oenanthe). Non-breeders established prospecting territories (<2 ha) that overlapped the breeding territories of conspecifics. We tested whether: (i) prospectors used social and environmental cues that predicted territory-specific breeding success in the following year, and (ii) the prospecting territory was tightly linked to the subsequent breeding territory of the prospector, and whether this link would be weakened by intraspecific competition with original territory owners if they also survived. 3. As expected, prospectors were attracted to a combination of site-specific cues that predicted future breeding success, i.e. short ground vegetation, a successfully breeding focal pair and successful close neighbours. 4. Prospecting behaviour was directly linked to the choice of the following year's breeding territory: 79% of surviving prospectors established a breeding territory at their prospecting site in the following year, with their breeding success being higher than other individuals of the same age. As predicted, fidelity to the prospected site was strongly dependent on whether the original territory owner of the same sex had died or moved. 5. Our findings suggest that the use of multiple cues reduces the negative impact of stochasticity on the reliability of social cues at small spatial scales (e.g. territories) and hence increases the probability of breeding success in the next year. Also, the use of conspecific attraction (i.e. the preference for breeding aggregations) is selectively advantageous because individuals are more likely to find a vacancy in an aggregation as compared to a solitary site. By extension, we hypothesize that species life-history traits may influence the spatial scale of prospecting behaviour and habitat selection strategies.