The form, magnitude and temporal dynamics of selection on phenotypic plasticity will fundamentally shape eco-evolutionary responses to environmental variation, but such attributes have not been fully conceptualized or quantified in nature. We provide a general framework that conceptualizes the dynamics of selection on phenotypic plasticity in labile dichotomous traits, which commonly shape behaviour and life history. Specifically, we highlight distinctions between selection on expressed plasticity and selection on resulting phenotypes, effects of phenotypic switches in opposite directions, and the full selection dynamics emerging across temporal sequences of environmental conditions. To enact this framework, we quantified selection on early-life plasticity in the ecologically critical trait of seasonal migration versus residence, by fitting a novel multi-state model to spatio-seasonal resighting data from 13 newly fledged cohorts of partially migratory European shags (Gulosus aristotelis). We demonstrate strong and consistent directional selection against early-life plasticity, manifested as substantially lower juvenile survival after phenotypic switches from resident to migrant, but not after reverse switches from migrant to resident. Yet, evident short-term costs translated into weaker and fluctuating selection on plasticity given sequences of phenotypes expressed throughout initial months. We thereby reveal how complex forms of selection against early-life plasticity can arise yet be rapidly attenuated in nature.
Life-history traits expressed in early life can exhibit considerable among-cohort variation, which could substantially affect population age-structure and dynamics if initial variation persists into later life-stages. Yet, initial among-cohort variation could be reinforced, rapidly dissipated, or else completely reshaped by dynamic combinations of age-specific phenotypic plasticity and selective disappearance acting within and among cohorts. However, such effects have not been comprehensively quantified for any trait, precluding full prediction of the form and implications of phenotypic dynamics, and emerging age-specific life-history variation, in varying environments. We provide a framework for conceptualising phenotypic change resulting from joint and interacting effects of cohort-specific and age-specific plasticity and selective disappearance. We implement this framework by quantifying overall early-life age-specific phenotypic change (or stasis), and dissecting underlying dynamics of plasticity and selection, for the ecologically critical life-history trait of seasonal migration versus residence. We achieve this by fitting multi-state models to extensive multi-year ring-resighting data from 9358 colour-ringed European shags (Gulosus aristotelis) from 11 cohorts in a partially migratory population. The overall cross-cohort mean proportion of migrants versus residents remained approximately constant across the four winters following fledging, implying no overall change in the degree of seasonal migration with age. This stasis was underlain by consistently high cross-year individual phenotypic repeatability, and by average plasticity towards residence that was counter-acted by average selective disappearance of sub-adult residents. However, these cross-cohort means obscured substantial among-cohort variation in the initial degree of partial migration, and in subsequent joint effects of plasticity and selective disappearance. Here, plasticity and selection were not systematically associated within or across cohorts or ages, but rather reinforced versus counter-acted each other at different times, thereby reshaping the pattern of among-cohort variation in partial migration across ages. These results demonstrate that an absence of overall age-specific change in a key life-history trait, seasonal migration versus residence, obscures substantial underlying variation in both early-life plasticity and selective disappearance, generating complex phenotypic dynamics within individual cohorts. Standard cross-cohort analyses may therefore inadequately predict future spatio-seasonal dynamics, since novel age-specific life-histories could readily emerge given changing environmental drivers of plasticity and selection.
Parasites are a fundamental component of wild animal populations, often inducing sub-lethal chronic effects that impact host fitness and demography. However, the factors determining variation in infection burden are often poorly understood in wild systems. Environmental conditions can determine exposure to infection and the resources required to respond, but exhibit strong temporal variation. As environmental conditions are predicted to become more variable, it is crucial to understand how these conditions shape burden to predict the downstream effects on host populations. Early-life conditions can shape responses to infection, potentially leading to delayed effects of environmental variation on fitness. The extent to which these are mediated by resources and later-life conditions remains unclear and may vary between the sexes, who often differ in exposure risk and resource requirements. Here, we examine how differences in hatching and breeding conditions influence parasite burden throughout life. We utilise data from a long-term population study of European shags (Gulosus aristotelis) on the Isle of May, Scotland, in which there is substantial variation in the timing of breeding within and between years, and nematode parasite burden can be measured in vivo using endoscopy. We show that adult parasite burden is influenced by seasonal and annual differences in current and early life conditions, but different patterns were observed in adult males and females. Burdens increased across the season in chicks and adult females but not in adult males. Instead, early life effects better explained burden in adult males, with those hatching later and in productive years displaying lower burdens. This suggests that early life may shape behaviour, immunity, or physiological development, impacting subsequent infection. Our findings reveal complex temporal effects on parasitism in species breeding in fluctuating environments. Incorporating seasonal and sex-specific responses to parasitism is crucial to understanding how predicted environmental shifts could impact disease dynamics.
In iteroparous, socially monogamous species, individuals vary in the extent of mate fidelity across breeding attempts, often with important fitness consequences. Numerous studies have demonstrated intrinsic drivers of mate fidelity, notably previous breeding success and parental age. Environmental conditions may also influence mate fidelity, and the habitat-mediated hypothesis predicts that fidelity will be lower when environmental conditions are poor. However, limited testing of this hypothesis has been undertaken in longitudinal studies of single populations. Furthermore, studies have mainly focused on environmental conditions during the breeding season, yet conditions prior to breeding may be important for mate fidelity because this is a critical period for pair bond formation. We investigated the effects of prebreeding environmental conditions (onshore wind component and sea surface temperature) on mate fidelity over a 20-year period in the socially monogamous, iteroparous, long-lived marine bird, the European shag, Gulosus aristotelis. Average fidelity rate varied three-to four-fold between years. Mate fidelity was affected by prebreeding environmental conditions, being lower when onshore winds were more prevalent and sea surface temperature was higher. However, mate fidelity was more strongly affected by intrinsic factors, with higher rates when breeding success in the previous attempt and population density were higher, and among older females and middle-aged males. We found that mate fidelity affected timing of breeding, with faithful pairs laying earlier, and early laying pairs bred more successfully, but there was no independent effect of mate fidelity on breeding success. Our results support the habitat-mediated hypothesis whereby prebreeding environmental conditions affect individual pairing decisions. Given environmental conditions are predicted to change globally, further investigation of their impact on aspects of social behaviour in a range of species is warranted. Crown Copyright (c) 2025 Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour. This is an open access article under the CC BY license (http://creativecommons.org/licenses/ by/4.0/).
In studies of avian populations accurate sex identification facilitates the study of sex-linked behaviour, ecology, social structure, decision-making, and life histories. To improve molecular sex identification protocols for the black-legged kittiwake (Laridae: Rissa tridactyla, hereafter ‘kittiwake’), primers previously developed for other bird species were realigned against the kittiwake CHD1 Z and W chromosomes to generate a species-specific primer-pair. Amplicons of 602 bp and 375 bp were produced using typical PCR and gel electrophoresis procedures. The modified primer-pair was validated on a test sample of 138 kittiwakes from four Atlantic breeding colonies sexed a priori using head-bill length.
Light-immersion data from geolocators can be used to estimate behavioural budgets and energetics in seabirds throughout the annual cycle. However, all methods used to categorise time spent in behaviours rely on assumptions that are difficult to validate. Additional data, such as pressure and temperature data from time-depth recorders (TDRs), can help to refine these assumptions. We explore the utility of previous methods to derive behavioural budgets from light-immersion data using a dataset from Common Guillemots Uria aalge, where individuals were equipped with both a TDR and a solar Global Location Sensor (GLS), also known as a ‘geolocator’. We compared behavioural allocations from previous methods to those derived when also using TDR data. Previous methods used light-immersion data to distinguish between time foraging, active, and resting on the water, but the addition of TDR data revealed that these activities resulted in similar light and immersion levels. It was also more difficult to differentiate between rest and flight using light-immersion data alone. However, by using insights gained from combined light-immersion and TDR data, we developed an improved method to assign behaviours using light-immersion data alone, and provide an adjusted equation to use these data to calculate energetics in Guillemots. We recommend using our approach when processing light-immersion data; however, if detailed activity budgets (particularly foraging information) are required, we recommend using higher resolution loggers, e.g. integrated light-immersion-temperature-pressure devices. Our findings are likely to be relevant for studies of other seabird species (particularly other auks) that dive and spend most of their time at sea during winter.
Identifying key areas of animal distribution using individual movement data is fundamental for conservation planning, threat mitigation, and spatial management. Methodologies which define these areas based on measures of high density and abundance may overlook spatial heterogeneity in behaviour-specific distributions. This is particularly relevant for behaviours that occur at lower densities but are associated with increased exposure to specific environmental threats. We used a dataset of 566 GPS tracked individuals and 14 colonies of a vulnerable species of seabird, the black-legged kittiwake (Rissa tridactyla), to compare two methods for delineating key areas. The first method applies kernel density estimates, based on 50
Emerging infectious diseases are of major concern to animal and human health. Recent emergence of high pathogenicity avian influenza virus (HPAIV) (H5N1 clade 2.3.4.4b) led to substantial global mortality across a range of host species. Co-occurring species showed marked differences in mortality, generating an urgent need for better epidemiological understanding within affected populations. We therefore tested for antibodies, indicative of previous exposure and recovery, and for active viral infection in apparently healthy individuals (n = 350) across five co-occurring seabird species on the Isle of May, Scotland, during 2023, following H5N1 HPAIV associated mortality in the preceding summer. Antibody prevalence to AIV subtypes varied substantially between species, ranging from 1.1% in European shags (Gulosus aristotelis) (to H5) to 78.7% in black-legged kittiwakes (Rissa tridactyla) (to H16 or both H13 and H16), and between 31 and 41% for three auk species (H5, H16 or both). At least 20.4% of auks had antibodies to an as yet unidentified subtype, suggesting further subtypes circulating in the population. We found low levels of active, but asymptomatic, AIV infection in individuals (1.6–4.5%), but excluded this as H5N1. Our results emphasise the importance of testing healthy individuals to understand the prevalence of co-circulating AIV subtypes in wild populations, and the potential for future reassortment events which could alter virus behaviour and impact.
Understanding the responses of seabirds to climate-induced variations in phenology and abundance of their prey is key to developing ecosystem-based fisheries management measures that benefit higher trophic levels. The match/mismatch hypothesis (MMH) emphasizes the need to consider synchrony in the seasonal cycles of predators and prey, while the match/mismatch/abundance hypothesis (MMAH) proposes that prey abundance may reinforce/compensate mismatch effects. This study considers the effects of both variations in seasonal availability and abundance of lesser sandeel Ammodytes marinus on hatching, fledging and breeding success of 5 seabird species: black-legged kittiwake Rissa tridactyla , Atlantic puffin Fratercula arctica , razorbill Alca torda , common guillemot Uria aalge and European shag Gulosus aristotelis . Consistent with MMH, temporal asynchrony between sandeel availability and seabird breeding schedules affected productivity in 4 species. The effects of trophic asynchrony were either reinforced or compensated by sandeel abundance for some species, supporting MMAH. Breeding success in the late-breeding kittiwake was high when conditions favoured both high sandeel abundance and temporal synchrony while the cost of asynchrony could be compensated by high sandeel abundance in the earlier-breeding puffin. Differential effects of sandeel abundance and trophic synchrony at different stages of the seabird breeding season suggest that distinct mechanisms are involved. The effects were most evident in the most sandeel-reliant seabirds. As further disruption of sandeel phenology and abundance is anticipated under the current climate crisis, the present study is an important step towards understanding bottom-up effects of environmental change on higher trophic levels.
To acquire or retain a higher quality breeding site, individuals may occupy sites outside the breeding season, with those investing more time and energy in this benefiting from improved breeding success. However, despite this benefit, the occupancy patterns of individuals may vary. Occupancy may be influenced by the distance individuals travel from breeding sites during the nonbreeding season; individuals nearer the colony may undertake occupancy earlier and more frequently than conspecifics because of shorter commuting distances from migration and foraging locations. Occupancy may also be energetically costly and affect how individuals are able to allocate their time to other key behaviours such as foraging. However, our understanding of how occupancy behaviour relates to an individual's distribution and ability to balance time and energy allocated to other behaviours is limited. Using data from a population of common guillemots, Uria aalge, a colonially breeding seabird, on the Isle of May, U.K., we investigated how nonbreeding occupancy of breeding sites is related to at-sea distribution, and how much energy and time individuals allocate to behaviours throughout the nonbreeding season We used bird-borne geolocators and time-depth recorders to record distribution and estimate time allocated to behaviours including occupancy, flight and foraging. Individuals that remained nearer to the colony before their first return then returned earlier and had shorter bouts of absence thereafter. Individuals also experienced a trade-off in the time spent in occupancy or foraging. Our data allowed us to estimate the increase in foraging efficiency required to offset the lost foraging time in individuals that occupied breeding sites. Overall, despite its known benefits, individuals varied in their timing and pattern of occupancy. We suggest that achieving consistently high breeding success, via nonbreeding season occupancy, may depend on an individual's distribution and ability to forage efficiently throughout the nonbreeding season. (c) 2024 The Authors. Published by Elsevier Ltd on behalf of The Association for the Study of Animal Behaviour.
Migration is a common trait among many animals allowing the exploitation of spatiotemporally variable resources. It often implies high energetic costs to cover large distances, for example between breeding and wintering grounds. For flying or swimming animals, the adequate use of winds and currents can help reduce the associated energetic costs. Migratory seabirds are good models because they dwell in habitats characterized by strong winds while undertaking very long migrations. We tested the hypothesis that seabirds migrate through areas with favourable winds. To that end, we used the SEATRACK dataset, a multi‐colony geolocator tracking dataset, for two North Atlantic seabirds with contrasting flight capabilities, the black‐legged kittiwake Rissa tridactyla and the Atlantic puffin Fratercula arctica , and wind data from the ERA5 climate reanalysis model. Both species had on average positive wind support during migration. Their main migratory routes were similar and followed seasonally prevailing winds. The general migratory movement had a loop‐shape at the scale of the North Atlantic, with an autumn route (southward) along the east coast of Greenland, and a spring route (northward) closer to the British Isles. While migrating, both species had higher wind support in spring than in autumn. Kittiwakes migrated farther and benefited from higher wind support than puffins on average. The variation in wind conditions encountered while migrating was linked to the geographical location of the colonies. Generally, northernmost colonies had a better wind support in autumn while the southernmost colonies had a better wind support in spring, with some exceptions. Our study helps understanding how the physical environment shapes animal migration, which is crucial to further predict how migrants will be impacted by ongoing environmental changes.
Knowledge of seabird distributions plays a key role in seabird conservation and sustainable marine management, underpinning efforts to designate protected areas or assess the impact of human developments. Technological advances in animal tracking devices increasingly allow researchers to acquire information on the movement of birds from specific colonies. Nevertheless, most seabird colonies have not been subject to such tracking and another means must be found to assess their likely foraging distribution. Consequently, foraging range data collated and summarized across other tracking studies has often been used to estimate species‐level foraging distances for use within applied settings. However, generic species‐specific foraging ranges must be used with caution because of the amount of variation in seabird foraging behaviour at both the individual and colony levels. Specifically, although current reviews of seabird foraging ranges provide summary estimates of maximum foraging range, they typically do not assess the extent of among‐colony or among‐individual variation around such estimates. To address this, we conducted a variance component analysis of the maximum distance reached from the breeding colony per foraging trip (foraging range) using multi‐colony tracking datasets to estimate the degree of between‐individual, between‐year and between‐colony variation in foraging range in four UK breeding seabirds (Black‐legged Kittiwake Rissa tridactyla , Common Guillemot Uria aalge , Razorbill Alca torda and European Shag Gulosus aristotelis ). We also provide updated estimates of typical foraging ranges for each species and quantify the influence of breeding stage and colony size. Overall, between‐colony variation was typically the largest variance component, explaining 20–30% of the observed variation in foraging range across the four species. Individual‐level variation was also relatively large among Shag. In Kittiwake, Guillemot and Shag, but not Razorbill, average foraging ranges were positively associated with colony size. In addition, Kittiwake and Razorbill travelled further during incubation than during chick‐rearing. More generally, our estimates of mean foraging ranges for each species were subject to a high degree of uncertainty, which should be incorporated into impact assessments carried out using such data.
BACKGROUND:Fidelity to a given foraging location or route may be beneficial when environmental conditions are predictable but costly if conditions deteriorate or become unpredictable. Understanding the magnitude of fidelity displayed by different species and the processes that drive or erode it is therefore vital for understanding how fidelity may shape the demographic consequences of anthropogenic change. In particular, understanding the information that individuals may use to adjust their fidelity will facilitate improved predictions of how fidelity may change as environments change and the extent to which it will buffer individuals against such changes.METHODS:We used movement data collected during the breeding season across eight years for common guillemots, Atlantic puffins, razorbills, and black-legged kittiwakes breeding on the Isle of May, Scotland to understand: (1) whether foraging site/route fidelity occurred within and between years, (2) whether the degree of fidelity between trips was predicted by personal foraging effort, and (3) whether different individuals made more similar trips when they overlapped in time at the colony prior to departure and/or when out at sea suggesting the use of the same local environmental cues or information on the decisions made by con- and heterospecifics.RESULTS:All species exhibited site and route fidelity both within- and between-years, and fidelity between trips in guillemots and razorbills was related to metrics of foraging effort, suggesting they adjust fidelity to their personal foraging experience. We also found evidence that individuals used local environmental cues of prey location or availability and/or information gained by observing conspecifics when choosing foraging routes, particularly in puffins, where trips of individuals that overlapped temporally at the colony or out at sea were more similar.CONCLUSIONS:The fidelity shown by these seabird species has the potential to put them at greater risk in the face of environmental change by driving individuals to continue using areas being degraded by anthropogenic pressures. However, our results suggest that individuals show some flexibility in their fidelity, which may promote resilience under environmental change. The benefits of this flexibility are likely to depend on numerous factors, including the rapidity and spatial scale of environmental change and the reliability of the information individuals use to choose foraging sites or routes, thus highlighting the need to better understand how organisms combine cues, prior experience, and other sources of information to make movement decisions.
Eco-evolutionary responses to environmentally induced selection fundamentally depend on magnitudes of genetic variation underlying traits that facilitate population persistence. Additive genetic variances and associated heritabilities can vary across environmental conditions, especially for labile phenotypic traits expressed through early life. However, short-term seasonal dynamics of genetic variances are rarely quantified in wild populations, precluding inference on eco-evolutionary outcomes in seasonally dynamic systems. This limitation applies to seasonal migration versus residence, constituting one key trait where rapid microevolution could rescue partially migratory populations from changing seasonal environments. We fitted novel quantitative genetic 'capture-recapture animal models' to multi-generational pedigree and year-round resighting data from 11 cohorts of European shags (Gulosus aristotelis), to estimate season-specific additive genetic variances in liabilities to migrate, and in resulting expression of migration, in juveniles' first autumn and winter. We demonstrate non-negligible genetic variation underlying early-life migration, with twice as large additive genetic variances and heritabilities in autumn than winter. Since early-life survival selection on migration typically occurs in winter, highest genetic variation and strongest selection are seasonally desynchronized. Our results reveal complex within- and among-year dynamics of early-life genetic and phenotypic variation, demonstrating that adequate inference of eco-evolutionary outcomes requires quantifying microevolutionary potential on appropriate scales and seasonal timeframes.
1. Population dynamic and eco-evolutionary responses to environmental variation and change fundamentally depend on combinations of within- and among-cohort variation in the phenotypic expression of key life-history traits, and on corresponding variation in selection on those traits. Specifically, in partially migratory populations, spatio-seasonal dynamics depend on the degree of adaptive phenotypic expression of seasonal migration versus residence, where more individuals migrate when selection favours migration. 2. Opportunity for adaptive (or, conversely, maladaptive) expression could be particularly substantial in early life, through the initial development of migration versus residence. However, within- and among-cohort dynamics of early-life migration, and of associated survival selection, have not been quantified in any system, preventing any inference on adaptive early-life expression. Such analyses have been precluded because data on seasonal movements and survival of sufficient young individuals, across multiple cohorts, have not been collected. 3. We undertook extensive year-round field resightings of 9359 colour-ringed juvenile European shags Gulosus aristotelis from 11 successive cohorts in a partially migratory population. We fitted Bayesian multi-state capture-mark-recapture models to quantify early-life variation in migration versus residence and associated survival across short temporal occasions through each cohort's first year from fledging, thereby quantifying the degree of adaptive phenotypic expression of migration within and across years. 4. All cohorts were substantially partially migratory, but the degree and timing of migration varied considerably within and among cohorts. Episodes of strong survival selection on migration versus residence occurred both on short timeframes within years, and cumulatively across entire first years, generating instances of instantaneous and cumulative net selection that would be obscured at coarser temporal resolutions. Further, the magnitude and direction of selection varied among years, generating strong fluctuating survival selection on early-life migration across cohorts, as rarely evidenced in nature. Yet, the degree of migration did not strongly covary with the direction of selection, indicating limited early-life adaptive phenotypic expression. 5. These results reveal how dynamic early-life expression of and selection on a key life-history trait, seasonal migration, can emerge across seasonal, annual, and multi-year timeframes, yet be substantially decoupled. This restricts the potential for adaptive phenotypic, microevolutionary, and population dynamic responses to changing seasonal environments.
Many industries are required to perform population viability analysis (PVA) during the consenting process for new developments to establish potential impacts on protected populations. However, these assessments rarely account for density-dependent regulation of demographic rates. Excluding density-dependent regulation from PVA-based impact assessments is often assumed to provide a maximum estimate of impact and therefore offer a precautionary approach to assessment. However, there is also concern that this practice may unnecessarily impede the development of important industries, such as offshore renewable energy. In this study, we assess density-dependent regulation of breeding success in 31 populations of seabird. We then quantify the strength and form of this regulation using eight different formulations. Finally, we use PVA to examine how each formulation influences the recreation of observed dynamics (i.e. model validation), as well as the predicted absolute and relative population response to an extrinsic threat (i.e. model projection). We found evidence of both negative (n = 3) and positive (n = 5) regulation of seabird breeding success. In populations exhibiting negative regulation, excluding density-dependent regulation from PVA-based impact assessment allowed uncontrolled population growth, such that model outcomes became biologically implausible. By contrast, in populations exhibiting positive regulation, excluding density-dependent regulation provided an appropriate reconstruction of observed dynamics, but population decline was underestimated in some populations. We find that multiple formulations of density dependence perform comparably at the detection, validation and projection stages of analysis. However, we tentatively recommend using a log-linear or Weibull distribution to describe density-dependent regulation of seabird breeding success in impact assessments to balance accuracy with caution. Finally, we show that relative PVA metrics of impact assessment cannot necessarily be used to overcome PVA misspecification by assuming density independence in positively regulated populations. Synthesis and applications: We suggest that a density-dependent approach when performing PVA-based assessments for seabird populations will prevent biologically unrealistic, unconstrained population growth and therefore ensure meaningful PVA metrics in populations experiencing negative regulation. It will also maintain a precautionary approach for populations experiencing positive regulation, crucial when estimating impacts for these more vulnerable populations. These conclusions have immediate international application within the consenting processes for marine industries. We suggest that a density-dependent approach when performing PVA-based assessments for seabird populations will prevent biologically unrealistic, unconstrained population growth and therefore ensure meaningful PVA metrics in populations experiencing negative regulation. It will also maintain a precautionary approach for populations experiencing positive regulation, crucial when estimating impacts for these more vulnerable populations. These conclusions have immediate international application within the consenting processes for marine industries.image
Demographic correlations are pervasive in wildlife populations and can represent important secondary drivers of population growth. Empirical evidence suggests that correlations are in general positive for long-lived species, however little is known about the degree of variation among populations in relation to local conditions. For three widely geographically separated Atlantic puffin populations (Fratercula arctica), we compared the relative importance of survival-reproduction correlations for two cross-season correlations, reflecting either effects of non-breeding season or breeding season conditions. Demographic rates and their correlations were estimated with an integrated population model, and their respective contributions to variation in population growth were calculated using a transient-LTRE. Demographic correlations were positive for all three populations, but their strength differed. By comparing three populations with geographically distinct foraging areas throughout the year, this study shows that demographic correlations are, in part, driven by environmental conditions, which impacts their population viability and vulnerability to environmental change.
Climate warming can reduce food resources for animal populations. In species exhibiting parental care, parental effort is a 'barometer' of changes in environmental conditions. A key issue is the extent to which variation in parental effort can buffer demographic rates against environmental change. Seabirds breed in large, dense colonies and globally are major predators of small fish that are often sensitive to ocean warming. We explored the causes and consequences of annual variation in parental effort as indicated by standardised checks of the proportions of chicks attended by both, one or neither parent, in a population of common guillemots Uria aalge over four decades during which there was marked variation in marine climate and chick diet. We predicted that, for parental effort to be an effective buffer, there would be a link between environmental conditions and parental effort, but not between parental effort and demographic rates. Environmental conditions influenced multiple aspects of the prey delivered by parents to their chicks with prey species, length and energy density all influenced by spring sea surface temperature (sSST) in the current and/or previous year. Overall, the mean annual daily energy intake of chicks declined significantly when sSST in the current year was higher. In accordance with our first prediction, we found that parental effort increased with sSST in the current and previous year. However, the increase was insufficient to maintain chick daily energy intake. In contrast to our second prediction, we found that increased parental effort had major demographic consequences such that growth rate and fledging success of chicks, and body mass and overwinter survival of breeding adults all decreased significantly. Common guillemot parents were unable to compensate effectively for temperature-mediated variation in feeding conditions through behavioural flexibility, resulting in immediate consequences for breeding population size because of lower adult survival and potentially longer-term impacts on recruitment because of lower productivity. These findings highlight that a critical issue for species' responses to future climate change will be the extent to which behavioural buffering can offer resilience to deteriorating environmental conditions.
The management of predator-prey conflicts can be a key aspect of species conservation. For management approaches to be effective, a robust understanding of the predator-prey relationship is needed, particularly when both predator and prey are species of conservation concern. On the Isle of May, Firth of Forth, Scotland, numbers of breeding Great Black-backed Gulls Larus marinus, a generalist predator, have been increasing since the 1980s, which has led to increasing numbers of sympatrically breeding Atlantic Puffins Fratercula arctica being predated during the breeding season. This may have consequences for species management on the Isle of May and impact assessments of offshore windfarms in the wider Firth of Forth area. We used population viability analysis to quantify under what predation pressure the Atlantic Puffin population may decline and become locally extinct over a three-generation period. The predation level empirically estimated in 2017 (1120 Puffins per year) was not sufficient to drive a decline in the Puffin population. Rather, an increase to approximately 3000 Puffins per year would be required to cause a population decline, and >4000 to drive the population to quasi-extinction within 66 years. We discuss the likelihood of such a scenario being reached on the Isle of May, and we recommend that where predator-prey conflicts occur, predation-driven mortality should be regularly quantified to inform conservation management and population viability analyses associated with impact assessments.