Geographically distinct reproductive schedules in a changing world: Costly implications in captive Stonechats

With progressively faster global change, shifts in phenology, and distributional ranges are reported for an increasing number of species. The success of organisms at coping with novel seasonal conditions depends on the mechanisms that determine their schedules. Species that rely on fixed schedules and those that time their activities by predictive cues may be particularly constrained in their ability to accommodate changes. The present study examines rigid scheduling and its implications for breeding in captivity in an avian model taxon, the Stonechat (Saxicola torquata). Within their extensive breeding range, Stonechats differ geographically in migratory behavior and reproduce and molt under a wide range of daylengths (10-17 h). Stonechats time their activities by programs that involve circannual rhythms and photoperiodism. The study reports reproductive timing of four taxa (central European, Irish, Siberian, and Kenyan), relates it to laydates in the field, and investigates modifying influences of housing conditions and of social context. Reproductive consequences of timing programs were then tested by crossbreeding of taxa with different schedules. The study revealed persistent, population-specific breeding windows in captivity. Resident Stonechats from equatorial Kenya synchronized their reproductive cycles with the European summer, presumably in response to local photoperiod, and bred at similar times as northern migrants. In all other taxa schedules matched those in the field, but were timed slightly earlier in captivity and advanced by indoor keeping conditions. Influences of social context were negligible. In pairs with clutches, testes regressed slightly later than in pairs without clutches, but presence of a mate per se had no influence on breeding cycles. Accordingly, crossbreeding Stonechats were predicted to have limited capacity to adjust schedules to those of their mates. This prediction was tested by crossbreeding of single-clutched Siberian long-distance migrants with multiple-clutched European short-distance migrants. Males and females of both taxa retained their characteristic breeding schedules, regardless of their mate's activities. This led to dramatic loss of reproductive success in the population with the longer breeding season, European Stonechats. Siberian Stonechats were unable to profit from the presence of a sexually active mate, but they suffered no disadvantage from crossbreeding. In a changing world, inherited timing programs may severely constrain responses to novel conditions, impose schedule-dependent, asymmetric costs of hybridization, and contribute to directional gene flow or to reproductive isolation.     

Carry-over effects from passage regions are more important than breeding climate in determining the breeding phenology and performance of three avian migrants of conservation concern

Long distance migrants are declining more rapidly than residents, with birds that breed in Europe and winter in tropical Africa providing particularly clear examples. Causal mechanisms may include climate change, but are poorly understood partly because carry-over effects from non-breeding ranges can influence breeding performance. Using long-term data spanning four decades we assess how climatic variation in migrants’ winter, passage and breeding ranges determine timing of breeding and reproductive success. We do so for three Afro-European avian migrants of regional conservation concern (redstart, spotted flycatcher and wood warbler). We find that carry-over effects from passage regions consistently had stronger impacts on breeding phenology than breeding climate. Warm Mediterranean passage conditions promoted earlier breeding in all species, and redstarts also bred earlier following higher Sahel rainfall. Warmer springs on the breeding grounds promoted slightly earlier breeding in redstart and wood warbler, but not spotted flycatcher. Carry-over effects also typically influenced breeding performance to a greater extent than weather on the breeding grounds. Greater rainfall in the Sahel increased redstart brood size, warmer Mediterranean passage conditions increased spotted flycatcher brood size and, to a lesser extent, the number of wood warbler fledglings. In contrast to the concern regarding climate change impacts on migrants’ breeding grounds we found no evidence that warmer temperatures on the breeding grounds were associated with reduced reproductive performance. We thus find that climatic variation on the non-breeding grounds, especially passage regions, typically influenced migrants’ breeding phenology and demography more strongly than equivalent variation on the breeding sites. Such carry-over effects should be considered when assessing the causes of migrants’ marked population declines.     

The endocrine basis of the reproductive cycle in the king penguin (Aptenodytes patagonicus)

What is the breeding frequency of the king penguin? More than 30 years of study and five major investigations, did not answer this question owing to the length (14 months) and the spreading (four months of laying) of reproductive events.
Following intensively during three consecutive years nearly 1000 tagged birds in a small colony, we obtained the individual history of a large part of the population. To these demographic and phenological studies, we have associated an endocrinological study validating blood samples and experiments in another colony.
The success or failure of a breeding cycle determines the timing of the subsequent breeding attempt. At Crozet Islands, the king penguin can rear one chick every two years at best: a successful breeding is followed by unsuccessful late breeding (or by non-breeding). So a large part of the population displays an annual periodicity of breeding attempts but with low individual breeding success.
The restarting of the reproductive cycle appears basically to be annual, the pituitary responding to the photoperiod. Steroid patterns present classical variations of level according to the reproductive stage. Prolactin (PRL) and luteinizing hormone (LH) are antagonists but, in this species, PRL remains high without external stimulation. To relay after failure in the field. we have experimentally to break down endogenous PRL. So, we have demonstrated that PRL has a key role in the reproductive cycle.
Paradoxically, the king penguin is an irregular breeder, quasi-biennial, although physiologically an annual breeder. In winter, there is not enough food at sea to rear the chick and during four months, the chick fasts. So a successful breeding cycle, which can be shortened to only nine months in captivity, is extended to more than a year for ecological reasons.     

The effect of migration strategy and food availability on White Stork Ciconia ciconia breeding success

In the mid 1970s, the breeding populations of the migrant White Stork Ciconia ciconia were close to extinction in the northeastern region of France (Alsace). A re-introduction project was implemented, resulting in the year-round settlement of some individuals in the region, which rely on additional food supplied by humans during the winter. Today, both resident and migrant birds breed in the same areas and take food from rubbish dumps and humans (farmers). The effects of these anthropogenic influences, altering Stork behaviour, on Stork reproductive success are not known. The aim of this study was to test the influence of bird status (resident vs. migrant) and food availability (control nests vs. nests that benefit from high food supply) on reproductive success. In control nests, the mean laying date was earlier in resident than in migrant White Storks. There was also a clear seasonal decline in clutch size. For all nests, the numbers of eggs and hatchlings were higher in resident birds than in migrants, which can be attributed to the earlier breeding of resident Storks. The large broods of resident birds showed a high mortality rate, leading to the same fledgling success (fledglings/hatchlings) and number of fledglings as in migrants. Fledgling success and the number of fledglings were higher for nests close to a reliable food supply. In summary, although resident birds can breed earlier and produce more eggs than migrants, we found no advantage in terms of number of fledglings. The higher mortality rate of chicks found in pairs with a large brood could be caused by the deterioration of their habitat. Thus, the year-round settlement of Storks may not present a biological advantage if the quality of their habitat is not guaranteed by the conservation of their grasslands.     

Behavioral and physiological conflicts in migrants: the transition between migration and breeding

Progression of the vernal migratory life history stage to breeding presents a number of apparent behavioral and physiological conflicts. Features that characterize the migratory stage include: high mobility, sociality, repetitive cycles of feeding (hyperphagia and fattening) and migratory flight. Breeding comprises: sedentary, territorial and reproductive behaviors, an initial decline in hyperphagia and reduction of fuel stores. Because morphology, physiology and behavior change, the transition between stages cannot be instantaneous. In many species development of the reproductive system actually occurs during migration, but in others gonadal development may not commence until later. This variation in degree of overlap of migration and reproductive functions is not well understood, but may be related to migratory distance and length of the breeding season, which tends to be shorter at higher latitudes and altitudes. In these habitats, migrants may arrive at their breeding sites to find unpredictable conditions that cannot support breeding. At this juncture, migrants may retreat to refugia and prolong maintenance of facultative migratory functions, termed arrival biology, until conditions improve sufficiently to initiate breeding. In this review, we focus on the Pacific races of the white-crowned sparrow, Zonototrichia leucophyrs, in which the entire spectrum of migratory strategies are represented from resident to long distance migrants and about which much is known. This species presents a unique view of the appearance and variations in arrival biology. Focusing on the juncture between migration and breeding, we discuss the diversity of responses of congeners to a spectrum of environmental conditions that favor survival and reproductive success.     

Breeding timed to maximize reproductive success for a migratory songbird: the importance of phenological asynchrony

Phenological advances and trophic mismatches are frequently reported ecological consequences of climate warming. Trophic mismatches occur when phenological responses to environmental conditions differ among trophic levels such that the timing of resource demand by consumers becomes decoupled from supply. We used 25 years of demographic measurements of a migratory songbird (the black‐throated blue warbler Setophaga caerulescens) to compare its breeding phenology to the phenology of both its caterpillar prey and the foliage on which caterpillars feed. Caterpillar biomass in this forest did not show a predictable seasonal pulse. Nest initiation by warblers in this northern hardwood forest was therefore not timed to coincide with a peak in food availability for nestlings. Nonetheless, timing of first clutches was strongly associated with spring leaf expansion (slope ± SE = 0.56 ± 0.08 days per day of change in leaf phenology, R² = 0.66). Warblers adjusted the timing of breeding to early springs mainly by shortening the interval between arrival and clutch initiation, but this likely has limits because recent early springs are approaching the relatively inflexible dates when birds arrive on the breeding grounds. Although the timing of first nests did not match 1:1 with leaf‐out phenology, the adjustments in breeding time maximized mean annual reproductive success. Nest predation had the greatest effect on annual reproductive success, but the ability of nesting warblers to appropriately track leaf phenology accounted for effects on annual reproductive success comparable to the influence of variation in caterpillar abundance and conspecific density. Nesting phenology in black‐throated blue warblers was generally well matched to the timing of leaf‐out, even though the match was not 1:1. Without measurements of reproductive success, these unequal phenological shifts might otherwise have been interpreted as having negative ecological consequences.     

Experimental food supplementation reveals habitat-dependent male reproductive investment in a migratory bird

Environmental factors can shape reproductive investment strategies and influence the variance in male mating success. Environmental effects on extrapair paternity have traditionally been ascribed to aspects of the social environment, such as breeding density and synchrony. However, social factors are often confounded with habitat quality and are challenging to disentangle. We used both natural variation in habitat quality and a food supplementation experiment to separate the effects of food availability—one key aspect of habitat quality—on extrapair paternity (EPP) and reproductive success in the black-throated blue warbler, Setophaga caerulescens. High natural food availability was associated with higher within-pair paternity (WPP) and fledging two broods late in the breeding season, but lower EPP. Food-supplemented males had higher WPP leading to higher reproductive success relative to controls, and when in low-quality habitat, food-supplemented males were more likely to fledge two broods but less likely to gain EPP. Our results demonstrate that food availability affects trade-offs in reproductive activities. When food constraints are reduced, males invest in WPP at the expense of EPP. These findings imply that environmental change could alter how individuals allocate their resources and affect the selective environment that drives variation in male mating success.     

Human-resource subsidies alter the dietary preferences of a mammalian top predator

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.     

The importance of food quantity and quality for reproductive performance in alpine water pipits (Anthus spinoletta)

Studies relating reproduction to food availability are usually restricted to food quantity, but ignore food quality and the effects of habitat structure on obtaining the food. This is particularly true for insectivorous birds. In this study we relate measures of reproductive success, time of reproduction and nestling size of water pipits (Anthus spinoletta) to biomass, taxonomic composition and nutritional content of available food, and to vegetation structure and distance to feeding sites. Clutch size was positively correlated with the proportion of grass at the feeding sites, which facilitates foraging. This suggests that water pipits adapt their clutch size to environmental conditions. Also, pipits started breeding earlier and produced more fledglings when abundant food and a large proportion of grass were available, probably because these conditions allow the birds to gain more energy in less time. The number of fledglings was positively correlated with the energy content of available food. No significant relationships were found between feeding conditions and nestling size or the time that nestlings took to fledge. This suggests that water pipits do not invest more in individual nestlings when food conditions are favourable but rather start breeding earlier and produce more young. Taxonomic composition and nutritional content of prey were not correlated with any of the reproductive parameters, indicating that profitability rather than quality of food affects reproductive success. Received: 31 May 1996/Accepted: 12 August 1996     

Habitat fragmentation influences nestling growth in Mediterranean blue and great tits

In patchy forest areas, the size of the forest patch where birds breed has a strong influence on their breeding success. However, the proximate effects contributing to lowering the breeding success in small forest patches remain unclear; and a shortage of crucial resources in those forest patches has been suggested to account in some degree for this failure. With the aim to further investigate this issue, we have monitored the breeding cycle of blue and great tits in three ‘large’ forest patches (ranging between 26.5 and 29.6 ha) and twelve ‘small’ forest patches (ranging between 1.1 and 2.1 ha) in a Mediterranean area in central Spain, during three years (2011–2013). We also recorded the nestling diet inside the nest-boxes with the aid of handy-cams. Only males significantly differed between forest patch size categories; being on average younger and with better body condition in small patches for great and blue tits respectively. Reproductive traits did not vary between forest patch size categories, but the body condition of blue tit nestlings and the size of great tit nestlings did, being significantly better and larger respectively in large forest patches. The recruitment rate of blue tit nestlings was also higher in large patches. Regarding nestling diet, blue tits did not differ but great tits did, delivering a larger amount of caterpillars in large forest patches. Most variation in the reproductive traits occurred between years, probably due to annual differences in environmental conditions. This study suggests that food supply could be limiting the breeding success of birds above all in small patches, but also in large patches under particular environmental conditions.     

Environmental Conditions during Breeding Modify the Strength of Mass-Dependent Carry-Over Effects in a Migratory Bird

In many animals, processes occurring in one season carry over to influence reproductive success and survival in future seasons. The strength of such carry-over effects is unlikely to be uniform across years, yet our understanding of the processes that are capable of modifying their strength remains limited. Here we show that female light-bellied Brent geese with higher body mass prior to spring migration successfully reared more offspring during breeding, but only in years where environmental conditions during breeding were favourable. In years of bad weather during breeding, all birds suffered reduced reproductive output irrespective of pre-migration mass. Our results suggest that the magnitude of reproductive benefits gained by maximising body stores to fuel breeding fluctuates markedly among years in concert with conditions during the breeding season, as does the degree to which carry-over effects are capable of driving variance in reproductive success among individuals. Therefore while carry-over effects have considerable power to drive fitness asymmetries among individuals, our ability to interpret these effects in terms of their implications for population dynamics is dependent on knowledge of fitness determinants occurring in subsequent seasons.      

Reduced parental effort in relation to laying date in house sparrows (Passer domesticus): a study under controlled conditions

It is known that breeding success in birds declines with an advance in the laying date. In this paper, breeding success diminished with laying date in a house sparrow (Passer domesticus) population breeding in captivity. The principal cause of the cline in breeding success with date was a decrease in the female feeding rate. The parental effort in females decreased with the advance of the laying date, although food access was ad libitum throughout the breeding season. Consequently, this cline in breeding success may be due to a reduced parental investment with the advance of the laying date or to exhaustion due to previous reproductive events in the season. Hypotheses usually used to explain the cline in breeding success with date can not explain results in this study.     

Can conditions experienced during migration limit the population levels of birds?

Populations of migratory birds are usually considered to be limited by conditions in breeding or wintering areas, but some might be limited by conditions encountered on migration. This could occur at stopover sites where competition for restricted food supplies can reduce subsequent survival or breeding success, or during the flights themselves, when adverse weather can occasionally kill large numbers of individuals. Competition for food could act in a density-dependent manner and help to regulate populations, whereas weather effects are more likely to act in a density-independent manner. The evidence for these views is explored in this paper. When preparing for migration, birds must normally obtain more food per day than usual, in order to accumulate the body reserves that fuel their flights. Birds often concentrate in large numbers at particular stopover sites, where food can become scarce, thus affecting migratory performance. Rates of weight gain, departure weights, and stopover durations often correlate with food supplies at stopover sites, sometimes influencing the subsequent survival and reproductive success of individuals, which can in turn affect subsequent breeding numbers. Many studies have provided evidence for interference and depletion competition at stopover sites, relatively few for migration conditions influencing the subsequent breeding or survival of individuals, and even fewer for effects on subsequent breeding numbers. Migrants in flight occasionally suffer substantial mortality in storms, especially over water, sometimes involving many thousands of birds at a time. Other mass mortalities have resulted from atypical ‘winter-like’ weather, occurring soon after the arrival of summer migrants in their breeding areas or just before their departure in autumn. Again, many thousands of birds at a time have been killed in such incidents, causing reductions of 30–90% in local breeding densities. In some bird species, migration-related events can at times have substantial effects on the year-to-year changes in breeding population levels. Nonetheless, the difficulties involved in investigating migrating birds at different points on their migration routes have so far limited the number of studies on the influence of events during migration periods on population levels.     

Seasonal patterns in reproductive success of temperate‐breeding birds: Experimental tests of the date and quality hypotheses

For organisms in seasonal environments, individuals that breed earlier in the season regularly attain higher fitness than their late‐breeding counterparts. Two primary hypotheses have been proposed to explain these patterns: The quality hypothesis contends that early breeders are of better phenotypic quality or breed on higher quality territories, whereas the date hypothesis predicts that seasonally declining reproductive success is a response to a seasonal deterioration in environmental quality. In birds, food availability is thought to drive deteriorating environmental conditions, but few experimental studies have demonstrated its importance while also controlling for parental quality. We tested predictions of the date hypothesis in tree swallows (Tachycineta bicolor) over two breeding seasons and in two locations within their breeding range in Canada. Nests were paired by clutch initiation date to control for parental quality, and we delayed the hatching date of one nest within each pair. Subsequently, brood sizes were manipulated to mimic changes in per capita food abundance, and we examined the effects of manipulations, as well as indices of environmental and parental quality, on nestling quality, fledging success, and return rates. Reduced reproductive success of late‐breeding individuals was causally related to a seasonal decline in environmental quality. Declining insect biomass and enlarged brood sizes resulted in nestlings that were lighter, in poorer body condition, structurally smaller, had shorter and slower growing flight feathers and were less likely to survive to fledge. Our results provide evidence for the importance of food resources in mediating seasonal declines in offspring quality and survival.     

Environmental conditions but not nest composition affect reproductive success in an urban bird

Adjusting the composition of their nests, breeding birds can influence the environmental conditions that eggs and offspring experience. Birds often use feathers to build nests, presumably due to their insulating properties. The amount of feathers in nests is often associated with increased nestling survival and body condition. However, it is unclear whether these putative beneficial effects of adding feathers to nests are relevant in a wide range of environmental conditions. Here, we combine data on weather conditions and feathers in nests (i.e., nest composition) to investigate their relative contribution to reproductive success in the Eurasian tree sparrow (Passer montanus). Specifically, we investigate whether the effect of weather conditions on breeding success is modulated by the amount of feathers added to the nest. We found a strong negative effect of rainfall on the number of nestlings that successfully fledged per breeding attempt, but this negative effect was not mitigated by the amount of feathers in nests. We also found that the amount of feathers in nests varied along the breeding season, with nests containing more feathers early in the breeding season, when temperatures were lower. Despite considerable variation in nest composition, our results do not suggest an important role of feathers in nests protecting eggs or nestling tree sparrows against fluctuations in environmental conditions.     

Reproductive responses of birds to experimental food supplementation: a meta-analysis

Introduction

Food availability is an important environmental cue for animals for deciding how much to invest in reproduction, and it ultimately affects population size. The importance of food limitation has been extensively studied in terrestrial vertebrate populations, especially in birds, by experimentally manipulating food supply. However, the factors explaining variation in reproductive decisions in response to food supplementation remain unclear. By performing meta-analyses, we aim to quantify the extent to which supplementary feeding affects several reproductive parameters in birds, and identify the key factors (life-history traits, behavioural factors, environmental factors, and experimental design) that can induce variation in laying date, clutch size and breeding success (i.e., number of fledglings produced) in response to food supplementation.

Results

Food supplementation produced variable but mostly positive effects across reproductive parameters in a total of 201 experiments from 82 independent studies. The outcomes of the food effect were modulated by environmental factors, e.g., laying dates advanced more towards low latitudes, and food supplementation appeared not to produce any obvious effect on bird reproduction when the background level of food abundance in the environment was high. Moreover, the increase in clutch size following food addition was more pronounced in birds that cache food, as compared to birds that do not. Supplementation timing was identified as a major cause of variation in breeding success responses. We also document the absence of a detectable food effect on clutch size and breeding success when the target species had poor access to the feed due to competitive interactions with other animals.

Conclusions

Our findings indicate that, from the pool of bird species and environments reviewed, extra food is allocated to immediate reproduction in most cases. Our results also support the view that bird species have evolved different life-history strategies to cope with environmental variability in food supply. However, we encourage more research at low latitudes to gain knowledge on how resource allocation in birds changes along a latitudinal gradient. Our results also emphasize the importance of developing experimental designs that minimise competition for the supplemented food and the risk of reproductive bottle-necks due to inappropriate supplementation timings.

     

Environmental effects on variation and covariation in reproductive traits of Western Bluebirds

Summary Environmental conditions can influence the expression and correlations of phenotypic traits. I studied phenotypic plasticity in reproductive traits of Western Bluebirds breeding in northern Arizona. Data collected over 4 years on two contrasting habitats identified significant spatial and temporal variation in bluebird reproduction. Clutch size was similar over different environmental conditions whereas timing of clutch initiation, percent fledging success, and frequency of second nest attempts were flexible. Correlations between traits varied widely—often changing sign—among samples from different years or habitats. Correlations of traits with reproductive success were also dependent on environmental conditions. Variation in traits reflected behavioral responses by nesting adults to differences in time for breeding and feeding conditions. Density of trees differed between habitats and had opposing effects on these environmental variables; breeding seasons were generally longer, but feeding rates to nestlings were lower on the more open habitat. Late Spring snows delayed reproduction and increased the importance of limited time for breeding; feeding conditions were more influential following a dry Spring. This and other studies illustrate that data on phenotypic plasticity are important when evaluating the ecological and evolutionary forces underlying life histories.     

Heavy and persistent rainfall leads to brood reduction and nest failure in a passerine bird

The process of reproduction is a sensitive period for bird offspring, since they are exposed to environmental conditions for several weeks. Within the long reproductive period, adult birds do not only have to maintain their own body condition, they also have to ensure nutritional needs of their nestlings. In the worst case, if weather conditions are very harsh, breeding is not successful and fails. This suggests that weather conditions might be an important driver of breeding success. Here, we studied the effect of weather conditions (temperature and precipitation) on the survival of 350 nests of great tits Parus major during nestling period in six years (2010–2015). We used traditional generalized linear mixed model (binomial response variable) and a dedicated survival-analysis program (MARK) to analyze the data. The two approaches allowed us to highlight and compare the effect of weather conditions on a fine (nestling survival) and coarse (nest fate) scale. Both methods showed that precipitation explained most of the variation in individual nestling and overall nest survival during the 15-day nestling period. Heavy and persistent rainfall did not only lead to brood reduction, it ultimately also led to losses of the entire brood. Likely causes were the negative effects of precipitation on thermoregulation, food availability and predation risk. However, while reduced food availability most likely might have led to brood reduction through selective individual nestling death, predation might have resulted in total nest failures.     

Free housing for declining populations: Optimizing the provision of artificial breeding structures

The improvement of habitat quality and reproductive success through the implementation of artificial breeding structures is one of the most widespread in situ conservation strategies applied to the recovery of declining wildlife populations. Several past studies have monitored the use of artificial breeding structures in the wild, but virtually none of them have investigated which demographic and environmental factors actually determine their effectiveness in facilitating reproductive success. Therefore, the aim of the present study was to identify those factors influencing breeding success in artificial structures. With this purpose we surveyed a declining population of a keystone species of Mediterranean ecosystems, namely the European wild rabbit. In Doñana National Park, we sampled during the course of two years the breeding success of wild rabbits from 47 artificial warrens, experimentally provided under different demographic and environmental conditions. In order to determine the relative importance of such factors, we applied an AIC model selection procedure to alternative biological hypothesis groups, postulating that breeding success in artificial structures would primarily depend on population density, food availability and/or predator limitation. We found that the efficiency of artificial warrens in facilitating rabbit reproductive success can vary, with an increase associated with greater pasture availability. Predator limitation by fencing also positively affected rabbit reproduction, but only in combination with high pasture availability. On their own, fencing treatment and population reinforcements had low and null effectiveness facilitating reproduction, respectively, as well as entailing significant economic and biological costs. The best way to improve breeding success in artificial structures, therefore, was through their strategic placement in potential high-quality habitats, i.e. with suitable pasture availability. We suggest that vegetation management can effectively increase carrying capacity and consequently the target species’ reproductive success, especially in fast breeding species like the European wild rabbit. The provision of artificial breeding structures for conservation purposes may significantly benefit from the elaboration of concurrent experimental studies similar to the present research, which can help establish useful guidelines for optimizing future conservation efforts.     

Mechanisms and reproductive consequences of breeding dispersal in a specialist predator under temporally varying food conditions

Individual variation in breeding dispersal has extensive ecological and evolutionary consequences, but the factors driving individual dispersal behaviour and their fitness consequences remain poorly understood. Our data on dispersal events of a rodent‐specialist predator, the Eurasian kestrel Falco tinnunculus, over 20 years in western Finland offers a unique opportunity to explore the mechanisms underlying breeding dispersal behaviour and its reproductive consequences in a wild bird population. Sex, age, body condition and previous breeding success affected breeding dispersal. Dispersal distances were longer in females than in males as well as longer in yearlings than in older individuals. Body condition was positively correlated to breeding dispersal distances, particularly for females. The lowest dispersal distances were recorded for intermediate brood sizes in the year preceding dispersal. Our results highlight sex‐ and environment‐specific consequences of breeding dispersal on reproductive performance. During increase phases of the three‐year vole cycles, males dispersing further had lower reproductive performance after dispersal, whereas in females, long breeding dispersal distances were associated with increased breeding success under all environmental conditions. These results suggest benefits associated to breeding dispersal in females, potentially related to large spatio‐temporal variation in main food abundance and intensity of intra‐specific competition. Breeding dispersal of males was costly during increasing food abundance, indicating the potential fitness benefits of environmental familiarity in this migratory species. Overall, our results indicate that both individual traits and environmental factors interact to shape breeding dispersal strategies in wide‐ranging predator populations under fluctuating food conditions.