Predation risk determines breeding territory choice in a Mediterranean cavity-nesting bird community

Non-direct effects of predation can be an important component of the total effect of predation, modulating animal population and community dynamics. The isolated effects of predation risk on the spatial organisation of the breeding bird community, however, remains poorly studied. We investigated whether an experimentally increased predation risk prior to reproduction affected breeding territory selection and subsequent reproductive strategies in three Mediterranean cavity-nesting birds, i.e., the little owl Athene noctua, European roller Coracias garrulus and scops owl Otus scops. We found that territories used the previous year were more likely to be re-occupied when they belonged to the safe treatment rather than to the risky treatment. The first choice of breeders of all three species was for safe territories over risky ones. When all breeding attempts in the season (i.e., final occupation) were considered, breeders also preferred safe to risky sites. In addition, little owls laid larger eggs in risky territories than in safe territories. Our study provides experimental evidence of a rapid preventive response of the three most abundant species in a cavity-nesting bird community to a short-term manipulation of predation risk. This response highlights the key role of the non-direct effects of predation in modulating avian community organisation.     

Predation on artificial bird nests in chaparral fragments

Summary The predation rate of artificial bird nests was measured in disturbed chaparral habitat fragments and at an unfragmented site in coastal San Diego County, California USA. Local extinctions of chaparral birds has been previously shown to occur in these fragments. The predation rate was highest at the unfragmented site. Among fragments, predation was higher at moderately disturbed than at highly disturbed sites. These results suggest that nest predator species diversity or density is reduced in disturbed chaparral fragments. Nest predation is probably not the most important cause of the observed loss of chaparral breeding bird diversity in these fragments.     

Predation risks suppress lifetime fitness in a wild mammal

Prey often reduce predation risk at the cost of lower resource intake. The cumulative effects of such tradeoffs can alter resource allocation, demography and evolutionary processes. We show how the accumulation of risk effects reduces the growth rate of wild North American porcupines Erethizon dorsatum, and simulate three evolutionary responses related to lifetime reproductive success. Individual porcupines experiencing predation risk from fishers Pekania pennanti grew slower and gave birth to fewer offspring. Simulations show that predation risk alone can lead to population declines, and that a female can replace herself by investing more energy into reproduction or adult survival; females that only invest energy in juvenile survival cannot. We show that the accumulation of predation risk can reduce lifetime reproductive success in natural ecosystems. Estimating the contribution of predation risk, and how evolutionary responses can mediate consequences associated with predation risk, is necessary to understand the evolution of predator–prey systems.     

Predation risk tradeoffs in prey: effects on energy and behaviour

The complexity of behavioural interactions in predator-prey systems has recently begun to capture trait-effects, or non-lethal effects, of predators on prey via induced behavioural changes. Non-lethal predation effects play crucial roles in shaping population and community dynamics, particularly by inducing changes to foraging, movement and reproductive behaviours of prey. Prey exhibit trade-offs in behaviours while minimizing predation risk. We use a novel evolutionary ecosystem simulation EcoSim to study such behavioural interactions and their effects on prey populations, thereby addressing the need for integrating multiple layers of complexity in behavioural ecology. EcoSim allows complex intra- and inter-specific interactions between behaviourally and genetically unique individuals called predators and prey, as well as complex predator-prey dynamics and coevolution in a tri-trophic and spatially heterogeneous world. We investigated the effects of predation risk on prey energy budgets and fitness. Results revealed that energy budgets, life history traits, allocation of energy to movements and fitness-related actions differed greatly between prey subjected to low-predation risk and high-predation risk. High-predation risk suppressed prey foraging activity, increased total movement and decreased reproduction relative to low-risk. We show that predation risk alone induces behavioural changes in prey which drastically affect population and community dynamics, and when interpreted within the evolutionary context of our simulation indicate that genetic changes accompanying coevolution have long-term effects on prey adaptability to the absence of predators.     

Morph-dependent resource acquisition and fitness in a polymorphic bird

Understanding genetic colour polymorphism has proved a major challenge, both in terms of the underlying genetic mechanisms and the evolutionarily forces maintaining such genetic variation. In this context, genetic differences in aggression or competitive-related traits may covary with the expression of alternative phenotypes, and affect the evolutionary stability and maintenance of colour polymorphisms. Genetic red and black head-colour morphs of the Gouldian finch (Erythrura gouldiae) co-occur in temporally and geographically stable frequencies in sympatric populations. Gouldian finches are obligate cavity-nesters with highly specific preferences for nest-site morphometry that directly affect reproductive success. Because intra- and interspecific competition for high quality nest-sites is prevalent, and fitness is directly related to nest-site quality, we investigated the relative access (and consequences for reproductive success) of alternative morphs to this critical limiting resource in the wild. Red males defended higher quality nest-sites, and overcame greater levels of nest-site competition against conspecifics and superior heterospecific competitors than black males. Red-headed males also produced more fledglings (especially with red-headed females) than black-headed males, independent of nest-site quality. Finally, the independent (positive) effect of nest-site quality on reproductive success was confirmed. Such competitive asymmetries are important to relative selection among coexisting morphs, and are likely to contribute to the maintenance of alternative sympatric colour-morphs in wild populations.     

Evolution of resident bird breeding phenology in a landscape with heterogeneous resource phenology and carryover effects

It is generally expected that, in environments with pronounced seasonal resource peaks, birds’ reproductive success will be maximised when nestlings’ peak food demand coincides with the timing of high food availability. However in certain birds that stay resident over winter, earlier breeding leads juveniles to join the winter flock earlier, which by the prior residence effect increases their success in breeding territory competition. This trade-off between reproduction and competition may explain why, in certain species, breeding phenology is earlier and asynchronous with the resource. This study extends a previous model of the evolution of breeding phenology in a single habitat type to a landscape with two habitat types: ‘early’ and ‘late’ resource phenology. The offspring’s natal habitat type has a carryover effect upon their competitive ability regardless of which habitat type they settle in to potentially breed. We find that, when the difference in resource phenology between habitats is small (weak carryover effect), breeding phenology in the late habitat evolves to occur earlier and more asynchronously than in the early habitat, to compensate for the competitive disadvantage to juveniles raised there. However if the difference is large (strong carryover effect), then the reproductive cost of earlier breeding outweighs the benefit of the compensation, so instead breeding phenology in the late habitat evolves to become more synchronous with the resource. Recruitment is generally asymmetric, from early to late habitat type. However if the early habitat is less frequent in the landscape or produces fewer offspring, then the asymmetry is reduced, and if there is some natal habitat-type fidelity, then recruitment can have an insular pattern, i.e. most recruits to each habitat type come from that same habitat type. We detail the different scenarios in which the different recruitment patterns are predicted, and we propose that they have implications for local adaptation.     

Genetic and environmental influences on natal dispersal distance in a resident bird species

We analyzed more than 1,600 dispersal events from two populations of a North American cooperatively breeding woodpecker species to determine what factors influence natal dispersal distance and whether distance traveled affects reproduction later in life. We found significant heritability of natal dispersal distance, in both males and females, indicating substantial additive genetic variance for this behavioral trait. Natal dispersal distance additionally was affected by social and ecological factors: individuals dispersing in their first year of life moved longer distances than those staying on their natal site as helpers for a prolonged time prior to dispersal, and increasing territory isolation led to longer dispersal distances. Successful dispersers incurred fitness costs, with lifetime fledgling production (in both sexes) and lifetime production of recruits to the breeding population (in females only) decreasing with increasing natal dispersal distance. We conclude that natal dispersal distance has a genetic basis but is modulated by environmental and social factors and that natal dispersal distance in this species is (currently) under selection.     

Seasonal matching of habitat quality and fitness in a migratory bird

When species occupy habitats that vary in quality, choice of habitat can be critical in determining individual fitness. In most migratory species, juveniles migrate independently of their parents and must therefore choose both breeding and winter habitats. Using a unique dataset of marked black-tailed godwits (Limosa limosa islandica) tracked throughout their migratory range, combined with analyses of stable carbon isotope ratios, we show that those individuals that occupy higher quality breeding sites also use higher quality winter sites. This seasonal matching can severely inflate inequalities in individual fitness. This population has expanded over the last century into poorer quality breeding and winter habitats and, across the whole population; individual birds tend to occupy either novel or traditional sites in both seasons. Winter and breeding season habitat selection are thus strongly linked throughout this population; these links have profound implications for a wide range of population and evolutionary processes. As adult godwits are highly philopatric, the initial choice of winter habitat by juveniles will be critical in determining future survival, timing of migration and breeding success.     

Lifetime fitness correlates of natal dispersal distance in a colonial bird

1. Obtaining empirical evidence of the consequences of dispersal distance on fitness is challenging in wild animals because long-term, unbiased data on reproduction, survival and movement are notoriously difficult to obtain. 2. Lifetime fitness correlates of natal dispersal distance were studied in an isolated population of the facultatively colonial lesser kestrel Falco naumanni (Fleischer) monitored during 8 years at north-eastern Spain, where most birds (83%) dispersed from their natal colony to settle at distances ranging from 112 m to 136.5 km. 3. Neither annual breeding success nor age at recruitment was affected by natal dispersal distance. However, a capture-mark-recapture analysis revealed that survival during the year following recruitment decreased exponentially with dispersal distance, with differences of up to 15% between philopatrics and long-distance dispersers. In subsequent years, it remained similar irrespective of the natal dispersal distance moved. These results did not seem to be biased by long-distance dispersers settling differentially in the periphery of the population (which could emigrate permanently and be considered dead in future occasions) or within-individual consistency in successive dispersal distances, so our results appear to reflect genuine survival differences between dispersal tactics. 4. Average lifetime fledgling production, average lifetime recruitment success and rate-sensitive individual fitness (λ(ind)) also decreased with the distance from the natal to the first-breeding colony, indicating that dispersal decisions early in life affecting immediate survival prospects may translate into long-term fitness costs. 5. Both survival and lifetime fitness models including continuous dispersal distances significantly improved the characterization of the effect on fitness compared with models considering dispersal as a discrete process (i.e. dispersal vs. philopatry at a colony level). 6. Long-distance dispersers were more likely to establish new colonies regardless of whether they recruited in the centre or the periphery of the population, revealing their important role in the colonization of unoccupied patches. Individuals experienced a higher probability of mortality in small and newly funded colonies, so lifetime fitness costs of dispersal seem to be explained by recruitment in sites where average quality is low because of high uncertainty in survival prospects.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

Covariation between bill colouration and fitness components in a nocturnal bird

Vividly coloured chromatic signals play a key role in social and sexual signalling in diurnal birds, but their role is considered negligible in favour of achromatic (i.e. white, pure grey and black colourations) signals in nocturnal species. Here we studied colour variation and potential signalling of the yellow bill – a trait functioning as sexual signal in diurnal raptors during the breeding season in females of a truly nocturnal raptor, the little owl Athene noctua. We found that yellow‐red chroma of the bill was highly variable between individuals and positively correlated with female fitness prospects (i.e. brood size at fledging). In addition, we found that females with brighter bills were larger in size and produced owlets with a higher mass at fledging. This study suggests that yellow bill colouration in female little owls may potentially play a role in sexual signalling and may constitute the first evidence of chromatic colour signalling in a nocturnal bird.     

Mechanisms and fitness effects of interspecific information use between migrant and resident birds

Interactions with potential competitors are an important componentof habitat quality. Due to the costs of coexistence with competitors,a breeding habitat selection strategy that avoids competitorsis expected to be favored. However, many migratory birds appearto gain benefits from an attraction to the presence of residentbirds, even though residents are assumed to be competitivelydominant. Thus far the mechanisms of this habitat selectionprocess, heterospecific attraction, are unknown, and the consequencesfor resident birds of migrant attraction remain untested. Throughheterospecific attraction, migrants may gain benefits if thedensity or territory location of residents positively reflectshabitat quality, and/or they gain benefits through increasedfrequency of social interactions with residents in foragingor predator detection. In this experiment, we examined the reciprocaleffects of spatial proximity on fitness-related traits in migrantpied flycatcher (Ficedula hypoleuca) and resident great tit(Parus major) by experimentally forcing them to breed eitheralone or in close proximity to each other. Surprisingly, greattits bore all the costs of coexistence while flycatchers wereunaffected, even gaining slight benefits. In concert with anearlier study, these results suggest that flycatchers use titsas information about good-quality nest-site locations whilebenefits from social interactions with tits are possible butless important. We suggest that utilizing interspecific socialinformation may be a common phenomenon between species sharingsimilar resource needs. Our results imply that the effects ofinterspecific information use can be asymmetric and may thereforehave implications for the patterns and consequences of speciescoexistence.     

Predation risk as a cost of reproduction

Predation risk as a cost of reproduction in animals has recently received increased empirical and theoretical attention. Higher risk may be associated with all stages of reproduction. Examples of evolutionary responses to this increased risk include habitat choice, duration and timing of display and copulation, changes in brightness of breeding coloration, and changes in life history traits such as age of reproduction and reproductive effort.     

The role of climate in limiting European resident bird populations

Abstract Aim To study densities of eight groups of resident forest bird populations across Europe to examine their association with abiotic (temperature, precipitation) factors. Location Europe. Methods Densities of residents (and migrant birds, which were used as a control group) were extracted from published breeding bird censuses. For each census location we obtained geographical co‐ordinates (latitude and longitude), temperature and precipitation variables describing both breeding and non‐breeding seasons. Resident densities were first examined separately in relation to co‐ordinates and principal component axes, which were extracted from temperature and precipitation variables. The relative impact of each explaining factor (co‐ordinates and principal component axis) on resident densities was checked by using residual examination and partial correlation. Results Densities of resident birds were in general negatively correlated with both latitude and longitude and temperature and precipitation factors, but latitude and temperature proved to be the strongest individual factors along which resident densities varied. The higher the latitude or the lower the temperatures, the lower were the densities of most resident birds. Partial correlation analysis suggested that of those two factors, latitude was the dominant one. Both the density of resident and migrant birds decreased towards the north, but the decrease of residents was steeper, and in the north they comprised only a small fraction of breeding bird numbers, whereas further south they commonly comprise half of the breeding bird numbers. Main conclusions The best explanatory factor for resident densities was latitude. For this reason, it was difficult to separate the relative effect of each individual factor, because latitude partly describes all the original variables (temperature and precipitation). The results suggested that precipitation was of less importance and that the interplay between latitude and ambient temperatures was not the sole factor behind the strong effect of latitude. We suggest that the coupled effect of harshening climate and decreasing amount of available energy with increasing latitude is limiting the population sizes of resident birds. We also propose that the effect of those factors is most apparent in northern Fennoscandia, where resident densities are suggested to be lower than would be expected on the basis of summer time carrying capacity.     

Age-independent telomere length predicts fitness in two bird species

Telomeres are dynamic DNA-protein structures that form protective caps at the ends of eukaryotic chromosomes. Although initial telomere length is partly genetically determined, subsequent accelerated telomere shortening has been linked to elevated levels of oxidative stress. Recent studies show that short telomere length alone is insufficient to induce cellular senescence; advanced attrition of these repetitive DNA sequences does, however, reflect ageing processes. Furthermore, telomeres vary widely in length between individuals of the same age, suggesting that individuals differ in their exposure or response to telomere-shortening stress factors. Here, we show that residual telomere length predicts fitness components in two phylogenetically distant bird species: longevity in sand martins, Riparia riparia, and lifetime reproductive success in dunlins, Calidris alpina. Our results therefore imply that individuals with longer than expected telomeres for their age are of higher quality.     

Predation risk predicts use of a novel habitat

Predator–prey interactions are often highly co‐evolved, with selection over time for prey with morphological and behavioral traits that minimize predation risk. Consequently, in many environments prey choose among potential habitats according to their refuge value. It is unclear, however, when presented with new habitats, if prey are able to evaluate the predation risk of these relative to familiar habitats and utilize these in accordance with their value. We tested whether, along the east coast of the USA, native mud crabs Panopeus herbstii utilize the non‐native alga Gracilaria vermiculophylla according to its relative refuge value. Experiments examining predation by blue crabs Callinectes sapidus on mud crabs revealed that the non‐native alga had an intermediate refuge value relative to native oysters, which were the most protective, and unvegetated sediment, which was the least. In subsequent choice experiments, mud crabs selected oysters over alga over unvegetated sediment, in accordance with habitat refuge values. Further, in field experiments, the use of Gracilaria by mud crabs was inversely related to the proximity of the alga to the preferred habitat type, oysters, and was reduced by the presence of a blue crab predator. Consequently, mud crabs are utilizing the non‐native alga Gracilaria in accordance with its intermediate refuge value. The relative refuge value of non‐native vs native habitat‐forming species may provide a baseline expectation against which to measure the speed of learning and opportunism in the response of native prey to novel protective habitats.     

捕食风险对小型哺乳动物行为、神经和内分泌的影响

长期进化过程中,猎物形成了对捕食者先天的敏感性。捕食风险对小型哺乳动物的影响主要包括行为的改变、神经环路的激活和内分泌的变化。行为的改变包括抑制运动活动、降低非防御行为以及改变生活环境。一些研究用c-fos免疫组化的方法测定了大鼠在暴露于猫的气味后大脑被激活的区域,主要包括中间杏仁核、下丘脑腹内侧区、背内侧区、前乳头状核和导水管周围灰质（periaqueductal gray）。猎物对捕食应激的反应还表现在一些内分泌指标的改变,一些激素如皮质酮（Corticoserone,CORT）和促肾上腺皮质激素（Adrenocorticotropic hormone,ACTH）的含量会显著的增加,同时在一些雄性动物中,睾丸激素的含量会有所降低。捕食者和猎物的关系近年来已经成为很多学科研究的有用工具。     

Predation of experimental nests is linked to local population dynamics in a fragmented bird population

Artificial nest experiments (ANEs) are widely used to obtain proxies of natural nest predation for testing a variety of hypotheses, from those dealing with variation in life-history strategies to those assessing the effects of habitat fragmentation on the persistence of bird populations. However, their applicability to real-world scenarios has been criticized owing to the many potential biases in comparing predation rates of artificial and natural nests. Here, we aimed to test the validity of estimates of ANEs using a novel approach. We related predation rates on artificial nests to population viability analyses in a songbird metapopulation as a way of predicting the real impact of predation events on the local populations studied. Predation intensity on artificial nests was negatively related to the species' annual population growth rate in small local populations, whereas the viability of large local populations did not seem to be influenced, even by high nest predation rates. The potential of extrapolation from ANEs to real-world scenarios is discussed, as these results suggest that artificial nest predation estimates may predict demographic processes in small structured populations.     

Preen oil and bird fitness: a critical review of the evidence

The uropygial gland is a holocrine complex exclusive to birds that produces an oleaginous secretion (preen oil) whose function is still debated. Herein, I examine critically the evidence for the many hypotheses of potential functions of this gland. The main conclusion is that our understanding of this gland is still in its infancy. Even for functions that are considered valid by most researchers, real evidence is scarce. Although it seems clear that preen oil contributes to plumage maintenance, we do not know whether this is due to a role in reducing mechanical abrasion or in reducing feather degradation by keratinophilic organisms. Evidence for a function against pathogenic bacteria is mixed, as preen oil has been demonstrated to act against bacteria in vitro, but not in vivo. Nor is it clear whether preen oil can combat pathogenic bacteria on eggshells to improve hatching success. Studies on the effect of preen oil against dermatophytes are very scarce and there is no evidence of a function against chewing lice. It seems clear, however, that preen oil improves waterproofing, but it is unclear whether this acts by creating a hydrophobic layer or simply by improving plumage structure. Several hypotheses proposed for the function of preen oil have been poorly studied, such as reduction of drag in flight. Similarly, we do not know whether preen oil functions as repellent against predators or parasites, makes birds unpalatable, or functions to camouflage birds with ambient odours. On the other hand, a growing body of work shows the important implications of volatiles in preen oil with regard to social communication in birds. Moreover, preen oil clearly alters plumage colouration. Finally, studies examining the impact of preen oil on fitness are lacking, and the costs or limitations of preen‐oil production also remain poorly known. The uropygial gland appears to have several non‐mutually exclusive functions in birds, and thus is likely to be subject to several selective pressures. Therefore, future studies should consider how the inevitable trade‐offs among different functions drive the evolution of uropygial gland secretions.     

Predation effects on the evolution of life-history traits in a clonal oligochaete

Although size at maturity and size and number of offspring are life-history traits widely studied in sexual and parthenogenetic reproduction, there is no such research on animals reproducing asexually without the involvement of gametes. Here we present an individual-based model in combination with experiments to study the clonal growth of Stylaria lacustris, an oligochaete reproducing through fission. We studied the effect of individual size at fission and fission ratio on clone fitness. Our results show that in benign environments without predators, fitness is higher when small worms produce small offspring. Then we included size-specific sublethal predation and found that the fitness of the clone is maximized when parental worms start fission at a large size and produce large descendants intercalated in the middle of the parental worm's body. These results agree with empirical findings. Furthermore, the results of our own laboratory experiment revealed that when S. lacustris is exposed to chemical alarm signals from injured conspecifics, it alters its life history in the same direction as predicted by the model. Our findings suggest that the effect of size-specific sublethal predation is similar to the effect of size-specific lethal predation because both modes of predation result in size-dependent prey mortality.