Long-distance migrants vary migratory behaviour as much as short-distance migrants: An individual-level comparison from a seabird species with diverse migration strategies.

As environmental conditions fluctuate across years, seasonal migrants must determine where and when to move without comprehensive knowledge of conditions beyond their current location. Animals can address this challenge by following cues in their local environment to vary behaviour in response to current conditions, or by moving based on learned or inherited experience of past conditions resulting in fixed behaviour across years. It is often claimed that long-distance migrants are more fixed in their migratory behaviour because as distance between breeding and wintering areas increases, reliability of cues to predict distant and future conditions decreases. While supported by some population-level studies, the influence of migration distance on behavioural variation is seldom examined on an individual level. Lesser black-backed gulls Larus fuscus are generalist seabirds that use a diversity of migration strategies. Using high-resolution multi-year GPS tracking data from 82 individuals from eight colonies in Western Europe, we quantified inter- and intra-individual variation in non-breeding distributions, winter site fidelity, migration routes and timing of migration, with the objectives of determining how much variation lesser black-backed gulls have in their migratory behaviour and examining whether variation changes with migration distance. We found that intra-individual variation was significantly lower than variation between individuals for non-breeding distributions, winter site fidelity, migration routes and timing of migration, resulting in consistent individual strategies for all behaviours examined. Yet, intra-individual variation ranged widely among individuals (e.g. winter site overlap: 0-0.91 out of 1; migration timing: 0-192 days), and importantly, individual differences in variation were not related to migration distance. The apparent preference for maintaining a consistent strategy, present in even the shortest distance migrants, suggests that familiarity may be more advantageous than exactly tracking current environmental conditions. Yet, variation in behaviour across years was observed in many individuals and could be substantial. This suggests that individuals, irrespective of migration distance, have the capacity to adjust to current conditions within the broad confines of their individual strategies, and occasionally, even change their strategy.

Are offspring begging levels exaggerated beyond the parental optimum? Evidence from a bidirectional selection experiment.

Parental care involves elaborate behavioural interactions between parents and their offspring, with offspring stimulating their parents via begging to provision resources. Thus, begging has direct fitness benefits as it enhances offspring growth and survival. It is nevertheless subject to a complex evolutionary trajectory, because begging may serve as a means for the offspring to manipulate parents in the context of evolutionary conflicts of interest. Furthermore, it has been hypothesized that begging is coadapted and potentially genetically correlated with parental care traits as a result of social selection. Further experiments on the causal processes that shape the evolution of begging are therefore essential. We applied bidirectional artificial selection on begging behaviour, using canaries (Serinus canaria) as a model species. We measured the response to selection, the consequences for offspring development, changes in parental care traits, here the rate of parental provisioning, as well as the effects on reproductive success. After three generations of selection, offspring differed in begging behaviour according to our artificial selection regime: nestlings of the high begging line begged significantly more than nestlings of the low begging line. Intriguingly, begging less benefitted the nestlings, as reflected by on average significantly higher growth rates, and increased reproductive success in terms of a higher number of fledglings in the low selected line. Begging could thus represent an exaggerated trait, possibly because parent-offspring conflict enhanced the selection on begging. We did not find evidence that we co-selected on parental provisioning, which may be due to the lack of power, but may also suggest that the evolution of begging is probably not constrained by a genetic correlation between parental provisioning and offspring begging.

Assay optimisation and age-related baseline variation in biochemical markers in Lesser Black-backed gulls.

Free-ranging animals are often used as bioindicators of both short- and long-term changes in ecosystem health, mainly to detect the presence and effects of contaminants. Birds, and gulls in particular, have been used as bioindicators over a broad range of marine and terrestrial ecosystems. In this study, we standardise the conditions for the use of a suite of biochemical markers in non-destructive matrices of Lesser Black-backed Gull (Larus fuscus) to facilitate future biomonitoring of marine and terrestrial contaminants. We characterized cholinesterase (ChE) in plasma and optimized assay conditions for ChE activity as a marker of neurotoxic damage. Moreover, we quantified variation in activity of ChE, lactate dehydrogenase (LDH), glutathione-S-transferase (GST) and catalase (CAT) as well as variation ranges of lipid peroxidation (LPO), in free-ranging adults and captive chicks. The main ChE form present in plasma of both adults and chicks was butyrylcholinesterase (BChE) followed by acetylcholinesterase (AChE), whose relative proportion in plasma tended to decrease with increased chick age. LPO levels and GST activity in blood cells (BCs) decreased significantly with increasing chick age, while BChE and LDH activity in plasma were not age-dependent. CAT in BCs tended to decline non-significantly in older chicks. Results of this study underscore the importance of standardising assay conditions and assessing intrinsic baseline variation in biochemical markers, before biochemical quantification. Data presented here provide a foundation for future use of BChE and LDH activity in plasma, as well as oxidative stress markers (LPO, CAT and GST) in BCs, to monitor environmental stress effects in Lesser Black-backed gulls.

Sex-specific effects of inbreeding on reproductive senescence.

Inbreeding depression plays a significant role in evolutionary biology and ecology. However, we lack a clear understanding of the fitness consequences of inbreeding depression. Studies often focus on short-term effects of inbreeding in juvenile offspring, whereas inbreeding depression in adult traits and the interplay between inbreeding depression and age are rarely addressed. Inbreeding depression may increase with age and accelerate the decline in reproductive output in ageing individuals (reproductive senescence), which could be subject to sex-specific dynamics. We test this hypothesis with a longitudinal experimental study in a short-lived songbird. Adult inbred and outbred male and female canaries were paired in a 2 × 2 factorial design, and survival and annual reproductive performance were studied for 3 years. We found inbreeding depression in female egg-laying ability, male fertilization success and survival of both sexes. Annual reproductive success of both males and females declined when paired with an inbred partner independent of their own inbreeding status. This shows that inbreeding can have fitness costs in outbred individuals when they mate with an inbred individual. Further, inbred females showed faster reproductive senescence than outbred females, confirming that inbreeding depression and age can interact to affect fitness. By contrast, there was no evidence for an interaction between inbreeding depression and reproductive senescence in male fertilization success. Our findings highlight the importance of considering sex-specific effects and age to determine the full range of fitness consequences of inbreeding and demonstrate that inbreeding depression can accelerate reproductive senescence.

Coadaptation of offspring begging and parental provisioning: A role for prenatal maternal effects?

Recent studies on birds have shown that offspring begging and parental provisioning covary at the phenotypic level, which is thought to reflect genetic correlations. However, prenatal maternal factors, like yolk testosterone, may also facilitate parent-offspring coadaptation via their effects on offspring begging and development. In fact, maternal effects are thought to adjust offspring phenotype to the environmental conditions they will experience after birth, which are in turn strongly dependent on the levels of parental provisioning. Using cross-fostering experiments in canaries, we tested the role of maternal effects on parent-offspring coadaptation from two different approaches. First, we analyzed whether females deposit yolk testosterone in relation to their own or their partner's prospective parental provisioning, measured as the rate of parental feeding to foster nestlings. Second, we investigated whether females deposit yolk testosterone in relation to costs they incurred when raising a previous brood, as this likely impinges on their capacity to provide parental care in the near future. However, from the results of both experiments we have no evidence that canary females deposit yolk testosterone in order to match offspring begging to the levels of care they and/or their partners provide. We therefore found no evidence that yolk testosterone facilitates parent-offspring coadaptation. In addition, our results suggest that the functional consequences of yolk testosterone deposition may relate to hatching asynchrony since it primarily varied with egg laying order.

'Out of tune': consequences of inbreeding on bird song.

The expression of bird song is expected to signal male quality to females. 'Quality' is determined by genetic and environmental factors, but, surprisingly, there is very limited evidence if and how genetic aspects of male quality are reflected in song. Here, we manipulated the genetic make-up of canaries (Serinus canaria) via inbreeding, and studied its effects upon song output, complexity, phonetics and, for the first time, song learning. To this end, we created weight-matched inbred and outbred pairs of male fledglings, which were subsequently exposed to the same tutor male during song learning. Inbreeding strongly affected syllable phonetics, but there were little or no effects on other song features. Nonetheless, females discriminated among inbred and outbred males, as they produced heavier clutches when mated with an outbred male. Our study highlights the importance of song phonetics, which has hitherto often been overlooked.

Influence of mate preference and laying order on maternal allocation in a monogamous parrot species with extreme hatching asynchrony.

It is well established that in many avian species, prenatal maternal resource allocation varies both between and within clutches and may affect offspring fitness. Differential allocation of maternal resources, in terms of egg weight and yolk composition, may therefore allow the female to adjust brood reduction and to fine-tune reproductive investment in accordance with the expected fitness returns. The adaptive value of such maternal resource allocation is thought to be context-dependent as well as species-specific. We investigated the effects of female preference for her mate on the allocation of prenatal maternal resources in the budgerigar, Melopsittacus undulatus, a monogamous species of parrot that shows an extreme hatching asynchrony. We assessed mate preferences in a two-way preference test and allowed females two breeding rounds: one with the preferred and one with the non-preferred partner. We found no effect of preference on either latency to lay or clutch size, but females mated with the preferred partner laid eggs that contained significantly more yolk. Their eggs also contained significantly more androstenedione but not testosterone. Our results suggest that in this species, female preference may influence maternal resource allocation, and that the functional roles of each androgen in the yolk should be considered separately. In addition, we found a significant effect of laying order on egg and yolk weight as well as on yolk testosterone and androstenedione levels. These measures, however, did not change linearly with the laying order and render it unlikely that female budgerigars compensate for the extreme hatching asynchrony by adjusting within-clutch allocation of prenatal maternal resources.

Testosterone influences song behaviour and social dominance - but independent of prenatal yolk testosterone exposure.

In the last two decades, maternally derived yolk androgens have been shown to significantly alter offspring development, and a number of these effects persist into adulthood. However, little is known about their underlying mechanisms. Mechanisms that have been suggested are changes in the endogenous androgen production post-hatching or changes in the sensitivity towards circulating androgens. We tested the effects of yolk testosterone on the plasma testosterone levels and the sensitivity to testosterone in 5months old male canaries that hatched from eggs that were either injected with testosterone (yT-males) or with a control solution (yC-males). Changes in sensitivity were investigated via the behavioural response to an experimental elevation of the plasma testosterone levels. We performed the experiment in fall (low endogenous testosterone production), focusing on testosterone dependent response traits (aggression and song). Before implantation, there was a non-significant trend that the plasma testosterone levels were lower in yT-males than in yC-males. Elevating the plasma testosterone concentrations increased aggressiveness, song bout length and similarity of repeated song elements (=consistency), with the latter likely being a consequence of testosterone-driven song crystallization. However, these effects were not different among yT- or yC-males in any of the parameters. Thus, our findings render it unlikely that changes in the sensitivity to testosterone post-hatching would form the main underlying mechanism of hormone-mediated maternal effects in birds. Further experiments are urgently needed in order to understand the nature of the phenotypic effects resulting from embryonic exposure to maternal yolk testosterone.

Seed preference is only weakly linked to seed-type-specific feeding performance in a songbird.

The dehusking of seeds by granivorous songbirds is a complex process that requires fast, coordinated and sensory-feedback-controlled movements of beak and tongue. Hence, efficient seed handling requires a high degree of sensorimotoric skill and behavioural flexibility, since seeds vary considerably in size, shape and husk structure. To deal with this variability, individuals might specialise on specific seed types, which could result in greater seed handling efficiency of the preferred seed type, but lower efficiency for other seed types. To test this, we assessed seed preferences of canaries (Serinus canaria) through food choice experiments and related these to data of feeding performance, seed handling skills and beak kinematics during feeding on small, spindle-shaped canary seeds and larger, spheroid-shaped hemp seeds. We found great variety in seed preferences among individuals: some had no clear preference, while others almost exclusively fed on hemp seeds, or even prioritized novel seed types (millet seed). Surprisingly, we only observed few and weak effects of seed preference on feeding efficiency. This suggests that either the ability to handle seeds efficiently can be readily applied across various seed types, or alternatively, it may indicate that achieving high levels of seed-specific handling skills does not require extensive practice.

Seed preference is only weakly linked to seed-type-specific feeding performance in a songbird.

The dehusking of seeds by granivorous songbirds is a complex process that requires fast, coordinated and sensory-feedback-controlled movements of beak and tongue. Hence, efficient seed handling requires a high degree of sensorimotoric skill and behavioural flexibility, since seeds vary considerably in size, shape and husk structure. To deal with this variability, individuals might specialise on specific seed types, which could result in greater seed handling efficiency of the preferred seed type, but lower efficiency for other seed types. To test this, we assessed seed preferences of canaries (Serinus canaria) through food choice experiments and related these to data of feeding performance, seed-handling skills and beak kinematics during feeding on small, spindle-shaped canary seeds and larger, spheroid-shaped hemp seeds. We found great variety in seed preferences among individuals: some had no clear preference, while others almost exclusively fed on hemp seeds, or even prioritized novel seed types (millet seed). Surprisingly, we only observed few and weak effects of seed preference on feeding efficiency. This suggests either that the ability to handle seeds efficiently can be readily applied across various seed types, or alternatively, that achieving high levels of seed-specific handling skills does not require extensive practice.

Leakage of plastics and other debris from landfills to a highly protected lake by wintering gulls.

GENERAL CONTEXT: Gulls ingest plastic and other litter while foraging in open landfills, because organic matter is mixed with other debris. Therefore, gulls are potential biovectors of plastic pollution into natural habitats, especially when they concentrate in wetlands for roosting. NOVELTY: We quantified, for the first time, the flow of plastic and other anthropogenic debris from open landfills to a natural lake via the movement of gulls. We focused on Fuente de Piedra, an inland closed-basin lake in Spain that is internationally important for biodiversity. METHODOLOGY: In 2022, we sampled gull pellets regurgitated in the lake by lesser black-backed gulls Larus fuscus that feed on landfills, as well as their faeces, then characterized and quantified debris particles of ≥0.5 mm. By combining GPS and census data from 2010 to 2022, together with plastic quantification based on FTIR-ATR analysis, we estimated the average annual deposition of plastic and other debris by the wintering gull population into the lake. MAIN RESULTS: 86 % of pellets contained plastics, and 94 % contained other debris such as glass and textiles. Polyethylene (54 %), polypropylene (11.5 %) and polystyrene (11.5 %) were the main plastic polymers. An estimated annual mean of 400 kg of plastics were moved by gulls into the lake. Only 1 % of plastic mass was imported in faeces. DISCUSSION: Incorporating the biovectoring role of birds can provide a more holistic view of the plastic cycle and waste management. Biovectoring is predictable in sites worldwide where gulls and other waterbirds feed in landfills and roost in wetlands. We discuss bird deterrence and other ways of mitigating debris leakage into aquatic ecosystems.

Artificial light at night impairs inhibitory control in a wild songbird.

Anthropogenic stressors, such as artificial light at night (ALAN), increasingly affect the sleep behaviour and physiology of wild birds, particularly in areas where human activity is prevalent. To understand the consequences of the resulting sleep deprivation, it is essential to investigate whether the effects of sleep deprivation on cognitive performance, observed in humans, also occur in birds. Here, we studied the impact of sleep deprivation, induced by intermittent exposure to ALAN, on inhibitory control, vigilance behaviour, and exploratory behaviour in great tits. Furthermore, we hypothesised that the effect of ALAN could depend on an individual's natural sleep duration and the timing of sleep. To achieve these goals, we measured emergence and entry times from/into the nest box in the wild, before capturing the great tits. In captivity, half of the birds were exposed to intermittent ALAN, and cognitive performance was assessed the following morning for all birds. ALAN-exposed birds were less successful on the detour reach task and when they started pecking at the test tube, they pecked more often. However, neither of the effects was related to the natural sleep duration or timing, in contrast to our hypothesis, and there were no differences between the ALAN-exposed and non-exposed group in vigilance and exploratory behaviour. Thus, even one night of exposure to ALAN can negatively affect cognitive performance in wild birds, possibly with negative effects on their performance and survival.

Offspring plumage coloration as a condition-dependent signal in the blue tit.

In many species, offspring display conspicuous coloration already early in life, even though they might be very vulnerable to predation at this stage. However, most attention has been drawn to the conspicuous plumage displayed by adult individuals in a sexual context, while other signaling functions have been explored much less. Here, we investigated whether the yellow breast plumage of blue tit (Cyanistes caeruleus) nestlings shows patterns of condition dependence and hence signals individual quality, as has been described for adult birds. During three consecutive breeding seasons, we, therefore, explored the association between nestling body mass and three color components of the yellow breast plumage (i.e., UV chroma, carotenoid chroma, and total brightness), considering both within and among nest effects. Variation in carotenoid chroma was not related to body mass. However, UV chroma and total brightness varied with body mass on an among-nest level, suggesting that they might signal aspects of genetic quality or parental rearing capacity. Interestingly, we also found a within-nest effect of body mass on total brightness, suggesting that this is a good candidate for a condition-dependent signal within the family. Thus, other family members could rely on brightness to adjust their behavioral strategies, such as feeding behavior in parents. Our study thus reveals that certain color components of the yellow breast plumage might signal different aspects of offspring quality, and they might have a correlated signaling value across life-history stages.

When parents play favorites: brood demand shapes parental preference for offspring UV color.

Parents might initially produce more offspring than they might be able to raise. However, when offspring demand exceeds their parents´ rearing capacity, parents might shift care towards the offspring which yield greater fitness returns to achieve their optimal brood size via brood reduction. Such favoritism could rely on offspring signaling traits if these inform parents about offspring quality and hence about the pay-offs of their investment. Here we investigated whether favoritism of blue tit (Cyanistes caeruleus) parents for an offspring signal (i.e., ultraviolet (UV) plumage coloration) varies with brood demand. To test this, we experimentally blocked the UV reflectance of yellow breast feathers in half of the nestlings of each brood, and then we sequentially performed two opposing brood size manipulations to vary nestling demand below or above parental rearing capacity. In reduced broods, nestlings begged overall less intensely and gained more body mass, supporting that parental rearing capacities sufficed to satisfy brood demand. Moreover, in reduced broods, UV-blocked nestlings (i.e., low-quality offspring) were fed and prey-tested more often. Yet, they begged more than control nestlings, suggesting that they were perhaps treated differently by other family members or which they may exploit parental preferences beyond actual need (at least in reduced nests). Parents flexibly shifted their feeding rate and favoritism in response to short-term changes in family size, as there was no parental preference for enlarged broods. Such flexible parental feeding rules may allow parents to gain the upper hand in parent-offspring conflict. However, we did not find evidence that parental favoritism facilitated brood reduction, at least in conditions where demand was temporally enhanced.

Environmental effects shape the maternal transfer of carotenoids and vitamin E to the yolk.

INTRODUCTION: Maternal effects occur when the phenotype of the offspring is influenced by the phenotype of the mother, which in turn depends on her heritable state as well as on influences from the current and past environmental conditions. All of these pathways may, therefore, form significant sources of variation in maternal effects. Here, we focused on the maternal transfer of carotenoids and vitamin E to the egg yolk, using canaries as a model species. Maternal yolk carotenoids and vitamin E are known to generate significant phenotypic variation in offspring, representing examples of maternal effects. We studied the intra-individual consistency in deposition patterns across two years and the mother-daughter resemblance across two generations in order to estimate the level of heritable variation. The effects of the current environmental conditions were studied via a food supplementation experiment, while the consequences of past environmental conditions were estimated on the basis of the early growth trajectories. RESULTS: There was a significant effect of the current environmental conditions on the yolk carotenoid and vitamin E deposition, but this effect varied between antioxidant components. The deposition of yolk carotenoids and vitamin E were linked to the process of yolk formation. Past environmental conditions did not contribute to the variation in yolk carotenoid and vitamin E levels nor did we find significant heritable variation. CONCLUSIONS: The transfer of carotenoids or vitamin E may be an example where current environmental variation is largely passed from the mother to the offspring, despite the numerous intermediate physiological steps that are involved. Differences in the effect of the environmental conditions as experienced by the mother during laying may be due to differences in availability as well as physiological processes such as competitive exclusion or selective absorption.

Sources of variation in yolk hormone deposition: consistency, inheritance and developmental effects.

Maternal effects occur when the phenotype of the mother affects the phenotype of their offspring. They are thought to have evolved to translate the environmental conditions experienced by the mother into adaptive phenotypic variation of the offspring. However, the integration of environmental cues allowing adaptive responses requires some form of plasticity that depends on the interaction of the maternal phenotype and her environment. In birds, maternal yolk hormones represent such a pathway for maternal effects, and their adaptive significance depends thus on the plasticity in maternal yolk hormone deposition. We studied sources of variation in yolk testosterone deposition, focusing on the often neglected contribution of the (partly heritable) maternal phenotype. We investigated consistency and heritability of yolk testosterone deposition in captive canaries of which the F(1) generation was raised in foster nests and analyzed the potential effects of the early developmental conditions. We found significant female consistency across years in egg mass, yolk mass and total amount of yolk testosterone but not in yolk testosterone concentrations. Females varied the yolk testosterone concentrations of their eggs across years mainly via changes in yolk mass. The heritable variation in egg mass, yolk mass and amount of yolk testosterone but not yolk testosterone concentrations was within the range of previous studies, but not significantly different from zero. Finally, the growth of the daughters as nestling had a significant effect on their yolk testosterone deposition at adulthood indicating the transgenerational potential for environmental effects - via the effects of yolk hormones on offspring development.

Food conditions affect yolk testosterone deposition but not incubation attendance.

In many bird species with hatching asynchrony, yolk androgens increase across the laying sequence. This has been hypothesized to represent a compensatory mechanism for disadvantages of later-hatching chicks - via positive effects of yolk androgens on early competitiveness and growth. However, the costs and benefits of this compensatory strategy probably depend on environmental factors determining the survival chances of the chicks such as the food conditions, which should, therefore, influence maternal yolk androgen deposition. We studied the consequences of manipulated food conditions on the expected level of hatching asynchrony in canaries (Serinus canaria) assigning females to either a low (=LQ) or high quality (=HQ) diet. We measured the incubation behaviour (as incubation attendance) and the yolk androgen deposition in order to investigate whether and how females modulate hatching asynchrony in relation to the food conditions. Females on a HQ diet laid larger and heavier clutches, showed a stronger increase in yolk testosterone content towards the last-laid eggs, but did not alter their incubation attendance. Thus, females on a HQ diet seem to favour the survival of later hatching chicks, as indicated by their yolk testosterone deposition pattern. However, females on a HQ diet laid larger clutches and might need to compensate more in order to achieve a similar degree of hatching asynchrony than females on a LQ diet, given the lack of plasticity in incubation attendance. This suggests that canary females respond to food manipulations mainly via changes in clutch size rather than by altering the degree of hatching asynchrony.

No relationship between chronotype and timing of breeding when variation in daily activity patterns across the breeding season is taken into account.

There is increasing evidence that individuals are consistent in the timing of their daily activities, and that individual variation in temporal behavior is related to the timing of reproduction. However, it remains unclear whether observed patterns relate to the timing of the onset of activity or whether an early onset of activity extends the time that is available for foraging. This may then again facilitate reproduction. Furthermore, the timing of activity onset and offset may vary across the breeding season, which may complicate studying the above-mentioned relationships. Here, we examined in a wild population of great tits (Parus major) whether an early clutch initiation date may be related to an early onset of activity and/or to longer active daylengths. We also investigated how these parameters are affected by the date of measurement. To test these hypotheses, we measured emergence and entry time from/into the nest box as proxies for activity onset and offset in females during the egg laying phase. We then determined active daylength. Both emergence time and active daylength were related to clutch initiation date. However, a more detailed analysis showed that the timing of activities with respect to sunrise and sunset varied throughout the breeding season both within and among individuals. The observed positive relationships are hence potentially statistical artifacts. After methodologically correcting for this date effect, by using data from the pre-egg laying phase, where all individuals were measured on the same days, neither of the relationships remained significant. Taking methodological pitfalls and temporal variation into account may hence be crucial for understanding the significance of chronotypes.

Intrinsic individual variation in daily activity onset and plastic responses on temporal but not spatial scales in female great tits.

In a variety of species, individuals appear to be consistent in the daily timing of their activity onset. Such consistent among-individual differences can result from both intrinsic factors, as individuals may e.g. differ genetically, and extrinsic factors, as the environment may vary on spatial and temporal scales. However, previous studies typically did not differentiate between their respective contributions on individual variation in the timing of activities. Here, we repeatedly measured the onset of activity in female great tits (Parus major) on consecutive days during the egg laying phase of the breeding season in four consecutive years. Subsequently, we used a variance partitioning analysis in order to determine which part of the total variation could be attributed to intrinsic (female identity) and extrinsic (nest box identity) factors. Overall, 27% of the total variation could be attributed to female identity. In addition, we found temporal variation in the activity onset, indicating that individuals can plastically adjust their timing. Yet despite their general ability to change the timing of activities over time, spatial environmental factors did not contribute significantly to the observed variation. Individuals may choose a habitat that matches the preferred timing of activities, or might not benefit from adjusting their timing to environmental factors that might vary on spatial scales.

Growing up in a crowd: social environment shapes the offspring's early exploratory phenotype in a colonial breeding species.

In colonial breeding species, the number of adverse social interactions during early life typically varies with breeding density. Phenotypic plasticity can help deal with this social context, by allowing offspring to adjust their behaviour. Furthermore, offspring may not be unprepared since mothers can allocate resources to their embryos that may pre-adjust them to the post-hatching conditions. Thus, we hypothesize that lesser black-backed gull chicks raised in dense breeding areas, with greater exposure to intra-specific aggression, show higher levels of anxiety and lower levels of exploration compared to chicks in low-density areas, and that this is facilitated by prenatal effects. To test this, we cross-fostered clutches within and across pre-defined high- and low-breeding density areas. We measured chicks' anxiety and exploration activity in an open-field test that included a novel and a familiar object. We found that both pre- and post-natal social environment contributed nearly equally and shaped the offspring's exploratory behaviour, but not its anxiety, in an additive way. Post-natal effects could reflect a learned avoidance of intra-specific aggression, yet identifying the pathways of the prenatal effects will require further study.