Experimentally quantifying the effect of nest-site depth on the predation risk and breeding success of Blue Tits

Predation is a key factor in the nesting preferences of birds. Studies indicate that cavity-breeding birds prefer deeper nest sites, possibly because they are more safe from predation. We studied the Blue Tit (Cyanistes caeruleus), a cavity-breeding passerine, to test (1) whether nest-site depth affects breeding success and (2) whether potential effects of nest-site depth on breeding success are related to predation risk. We performed 2 experiments to separate effects of nest-box depth from potential effects of the quality of the breeding pair. In the first (free-choice) experiment, Blue Tits competed for scarce deep nest boxes that were provided well before nest-box choice, enabling an association between nest-box quality and bird quality. In the second (forced-choice) experiment, we randomly altered nest-box depth after Blue Tits had chosen a nest box, thus disconnecting the association between nest-box quality and bird quality. We found no evidence that the occurrence of signs of predation was related to nest-box depth. However, we did find clear positive effects of nest-box depth (1) on clutch size and hatching success throughout the study area and (2) on fledging success, the number of fledglings, and the overall probability of nest success, specifically in parts of the study area with high predation. We found no indication of independent effects of parental quality on breeding success. Parents also seemed to perceive the shallower boxes as more risky; in shallower boxes, nest thickness was decreased, irrespective of the local predation pressure during the free-choice experiment. Parents nesting in shallow boxes may have had lower breeding success because of (1) increased actual (but undetected) predation and (2) reduced reproductive investment by parents, based on the latter's experience with predation or an evolutionary response to past predation risk

State dependence explains individual variation in nest defence behaviour in a long-lived bird

Parental care, such as nest or offspring defence, is crucial for offspring survival in many species. Yet, despite its obvious fitness benefits, the level of defence can consistently vary between individuals of the same species. One prominent adaptive explanation for consistent individual differences in behaviours involves state dependency: relatively stable differences in individual state should lead to the emergence of repeatable behavioural variation whereas changes in state should lead to a readjustment of behaviour. Therefore, empirical testing of adaptive state dependence requires longitudinal data where behaviour and state of individuals of the same population are repeatedly measured. Here, we test if variation in states predicts nest defence behaviour (a 'risky' behaviour) in a long-lived species, the barnacle goose Branta leucopsis. Adaptive models have predicted that an individual's residual reproductive value or 'asset' is an important state variable underlying variation in risk-taking behaviour. Hence, we investigate how nest defence varies as a function of time of the season and individual age, two state variables that can vary between and within individuals and determine asset. Repeated measures of nest defence towards a human intruder (flight initiation distance or FID) of females of known age were collected during 15 breeding seasons. Increasing values of FID represent increasing shyness. We found that females strongly and consistently differed in FID within- and between-years. As predicted by theory, females adjusted their behaviour to state by decreasing their FID with season and age. Decomposing these population patterns into within- and between-individual effects showed that the state-dependent change in FID was driven by individual plasticity in FID and that bolder females were more plastic than shyer females. This study shows that nest defence behaviour differs consistently among individuals and is adjusted to individual state in a direction predicted by adaptive personality theory

Reproductive effort and future parental competitive ability:A nest box removal experiment

The life history trade-off between current and future reproduction is a theoretically well-established concept. However, empirical evidence for the occurrence of a fitness cost of reproduction is mixed. Evidence indicates that parents only pay a cost of reproduction when local competition is high. In line with this, recent experimental work on a small passerine bird, the Great tit (Parus major) showed that reproductive effort negatively affected the competitive ability of parents, estimated through competition for high quality breeding sites in spring. In the current study, we further investigate the negative causal relationship between reproductive effort and future parental competitive ability, with the aim to quantify the consequences for parental fitness, when breeding sites are scarce. To this end, we (a) manipulated the family size of Great tit parents and (b) induced severe competition for nest boxes among the parents just before the following breeding season by means of a large-scale nest box removal experiment. Parents increased their feeding effort in response to our family size manipulation and we successfully induced competition among the parents the following spring. Against our expectation, we found no effect of last season's family size on the ability of parents to secure a scarce nest box for breeding. In previous years, if detected, the survival cost of reproduction was always paid after midwinter. In this year, parents did pay a survival cost of reproduction before midwinter and thus before the onset of the experiment in early spring. Winter food availability during our study year was exceptionally low, and thus, competition in early winter may have been extraordinarily high. We hypothesize that differences in parental competitive ability due to their previous reproductive effort might have played a role, but before the onset of our experiment and resulted in the payment of the survival cost of reproduction

Great tits trade off future competitive advantage for current reproduction

Quantifying the fitness costs of reproduction is essential to understand the evolution of reproductive behavior. Recent work shows that increased reproductive investment reduced parental survival in more competitive environments. Here we experimentally test the hypothesis that reproductive investment has a negative effect on the ability of parents to compete for resources in later life. In a nest-box population of Great tits (Parus major), we manipulated family size by reducing or enlarging broods with 2 or 3 nestlings, relative to a control group, in 2 years (N = 237 broods). Parental feeding effort was positively related to experimental family size but leveled off for the enlarged broods. In the next breeding season we manipulated the nest box quality in early spring by reducing the depth of 80% of the nest boxes (deeper boxes are safer and preferred). We analyzed parents' probability of obtaining a deep breeding box in relation to their previous year's family size manipulation. We found for both years that increased reproductive investment negatively affected the probability of parents to claim a high-quality nest box in the subsequent breeding season. We thus confirm that family size has a negative effect on the future competitive ability of parents. Such carry-over effects are important because they show that selection on individual optimal clutch size will depend on 1) resource abundance and the level of competition in the next breeding season and 2) the reproductive investment of the competitors in the current breeding season because it affects their future competitive ability as well

Social competition as a driver of phenotype–environment correlations: implications for ecology and evolution

Fokkema RW, Korsten P, Schmoll T, Wilson AJ. Social competition as a driver of phenotype–environment correlations: implications for ecology and evolution. Biological Reviews. 2021: brv.12768.While it is universally recognised that environmental factors can cause phenotypic trait variation via phenotypic plasticity, the extent to which causal processes operate in the reverse direction has received less consideration. In fact individuals are often active agents in determining the environments, and hence the selective regimes, they experience. There are several important mechanisms by which this can occur, including habitat selection and niche construction, that are expected to result in phenotype–environment correlations (i.e. non-random assortment of phenotypes across heterogeneous environments). Here we highlight an additional mechanism – intraspecific competition for preferred environments – that may be widespread, and has implications for phenotypic evolution that are currently underappreciated. Under this mechanism, variation among individuals in traits determining their competitive ability leads to phenotype–environment correlation; more competitive phenotypes are able to acquire better patches. Based on a concise review of the empirical evidence we argue that competition-induced phenotype–environment correlations are likely to be common in natural populations before highlighting the major implications of this for studies of natural selection and microevolution. We focus particularly on two central issues. First, competition-induced phenotype–environment correlation leads to the expectation that positive feedback loops will amplify phenotypic and fitness variation among competing individuals. As a result of being able to acquire a better environment, winners gain more resources and even better phenotypes – at the expense of losers. The distinction between individual quality and environmental quality that is commonly made by researchers in evolutionary ecology thus becomes untenable. Second, if differences among individuals in competitive ability are underpinned by heritable traits, competition results in both genotype–environment correlations and an expectation of indirect genetic effects (IGEs) on resource-dependent life-history traits. Theory tells us that these IGEs will act as (partial) constraints, reducing the amount of genetic variance available to facilitate evolutionary adaptation. Failure to recognise this will lead to systematic overestimation of the adaptive potential of populations. To understand the importance of these issues for ecological and evolutionary processes in natural populations we therefore need to identify and quantify competition-induced phenotype–environment correlations in our study systems. We conclude that both fundamental and applied research will benefit from an improved understanding of when and how social competition causes non-random distribution of phenotypes, and genotypes, across heterogeneous environments

Preen oil composition of Pied Flycatchers is similar between partners but differs between sexes and breeding stages

Preen oil, the secretion of the uropygial gland, may be an important source of body odour in birds. By characterizing the chemical composition of preen oil, we can describe the olfactory phenotypes of birds and investigate whether odours could have a function in sexual signalling or other chemical communication. Here we analysed the preen oil of a wild passerine, the European Pied Flycatcher Ficedula hypoleuca, to find out whether it holds socially relevant information. We sampled both the female and male of breeding pairs during nestling rearing to test for sex differences and within-pair similarity. We additionally sampled the females during incubation to test for changes across breeding stages and for individual repeatability of chemical profiles. Pair mates had similar chemical profiles in comparison with other breeding adults. Furthermore, we found evidence for sex differences and for changes across breeding stages. Notably, the preen oil of females was more diverse and more volatile than that of males, and the preen oil secreted by females during incubation was more volatile than that secreted during nestling rearing. However, we found no evidence for individual repeatability of chemical profiles across breeding stages in females. Our results point towards a function of preen oil in sexual signalling, although other functions should not be excluded. Our study is a first step towards understanding the role of odours in the social life of an important avian model species used in the study of mate choice and sexual selection.</p

Data from: Reproductive effort and future parental competitive ability: a nest box removal experiment

The life history trade-off between current- and future reproduction is a theoretically well-established concept. However, empirical evidence for the occurrence of a fitness cost of reproduction is mixed. Evidence indicates that parents only pay a cost of reproduction when local competition is high. In line with this, recent experimental work on a small passerine bird, the Great tit (Parus major) showed that reproductive effort negatively affected the competitive ability of parents, estimated through competition for high quality breeding sites in spring. In the current study we further investigate the negative causal relationship between reproductive effort and future parental competitive ability, with the aim to quantify the consequences for parental fitness, when breeding sites are scarce. To this end, we 1) manipulated the family size of Great tit parents and 2) induced severe competition for nest boxes among the parents just before the following breeding season by means of a large scale nest box removal experiment. Parents increased their feeding effort in response to our family size manipulation and we successfully induced competition among the parents the following spring. Against our expectation, we found no effect of last season’s family size on the ability of parents to secure a scarce nest box for breeding. In previous years, if detected, the survival cost of reproduction was always paid after midwinter. In this year parents did pay a survival cost of reproduction before midwinter and thus before the onset of the experiment in early spring. Winter food availability during our study year was exceptionally low and thus competition in early winter may have been extraordinarily high. We hypothesize that differences in parental competitive ability due to their previous reproductive effort might have played a role, but before the onset of our experiment and resulted in the payment of the survival cost of reproduction

Fokkema_et_al_2018_data_reprod_effort_and_comp_ability

The datafile is an excel file containing 5 sheets with the data used for the article. Please read the accompanying text file 'README.txt' for more information on the datasets and the meaning of the columns

Data from: A novel mechanism for grazing lawn formation: large herbivore-induced modification of the plant-soil water balance

1. Large herbivores play a key-role in creating spatial heterogeneity through the formation of grazing lawns. Recent research suggests that the currently accepted nutrient-based theory on the formation of these grazing lawns cannot universally explain their formation in all ecosystems where they are found. 2. We developed and investigated an alternative hypothesis on grazing lawn formation and maintenance based on herbivore effects on the plant-soil water balance. We propose that large herbivores change the soil water balance in grazing lawns through defoliation and soil compaction, causing a shift in vegetation composition towards a drought tolerant plant community. 3. Investigating this idea in a tropical savanna, we indeed found profound differences in grazing lawn soil properties and water balance. In particular, defoliation increased soil temperatures and potential evaporation rates while soil compaction increased bulk density and decreased water infiltration rates, especially on fine-textured soils. Soil moisture was therefore generally much lower in grazing lawns than in adjacent bunch grass areas. 4. Furthermore, we found that grazing lawn species show drought-tolerant traits, with higher leaf sodium levels, suggesting evolutionary adaptation to these herbivore-induced dry conditions. However, leaf water potentials did not differ between grazing lawn and bunch grass species. 5. Synthesis: This study shows that large herbivores might form grazing lawns through previously underestimated effects on water balance. Thus, future studies on large herbivore effects on vegetation should increasingly focus on additional pathways of soil compaction and defoliation. While nutrient-based processes driving grazing lawn formation may operate during the wet season in savannas, we suggest that water balance-based processes are additionally important during the dry season