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Migration is a widespread strategy for escaping unfavourable conditions during winter, but the extent to which populations that segregate during the breeding season aggregate during the non-breeding season is poorly understood. Low non-breeding season aggregation may be associated with higher likelihood of overlap with threats, but with fewer populations affected, whereas high aggregation may result in a lower probability of exposure to threats, but higher overall severity. We investigated non-breeding distributions and extent of population aggregation in 2 sympatrically breeding auks. We deployed geolocation-immersion loggers on common guillemots Uria aalge and razorbills Alca torda at 11 colonies around the northern UK and tracked their movements across 2 non-breeding seasons (2017-18 and 2018-19). Using 290 guillemot and 135 razorbill tracks, we mapped population distributions of each species and compared population aggregation during key periods of the non-breeding season (post-breeding moult and mid-winter), observing clear interspecific differences. Razorbills were largely distributed in the North Sea, whereas guillemot distributions were spread throughout Scottish coastal waters and the North, Norwegian and Barents Seas. We found high levels of aggregation in razorbills and a strong tendency for colony-specific distributions in guillemots. Therefore, razorbills are predicted to have a lower likelihood of exposure to marine threats, but more severe potential impact due to the larger number of colonies affected. This interspecific difference may result in divergent population trajectories, despite the species sharing protection at their breeding sites. We highlight the importance of taking whole-year distributions into account in spatial planning to adequately protect migratory species.

Capsule: A data-thinning approach was used to assess the effects of reducing the frequency of nest-checks on estimates of breeding success of Common Guillemots Uria aalge. Inter-year and inter-colo...

Understanding how ecological processes combine to shape population dynamics is crucial in a rapidly changing world. Evidence has been emerging for how fundamental drivers of density dependence in mobile species are related to two differing types of environmental variation-temporal variation in climate, and spatiotemporal variation in food resources. However, to date, tests of these hypotheses have been largely restricted to mid-trophic species in terrestrial environments and thus their general applicability remains unknown. We tested if these same processes can be identified in marine upper trophic level species. We assembled a multi-decadal data set on population abundance of 10 species of colonial seabirds comprising a large component of the UK breeding seabird biomass, and covering diverse phylogenies, life histories and foraging behaviours. We tested for evidence of density dependence in population growth rates using discrete time state-space population models fit to long time-series of observations of abundance at seabird breeding colonies. We then assessed if the strength of density dependence in population growth rates was exacerbated by temporal variation in climate (sea temperature and swell height), and attenuated by spatiotemporal variation in prey resources (productivity and tidal fronts). The majority of species showed patterns consistent with temporal variation in climate acting to strengthen density dependent feedbacks to population growth. However, fewer species showed evidence for a weakening of density dependence with increasing spatiotemporal variation in prey resources. Our findings extend this emerging theory for how different sources of environmental variation may shape the dynamics and regulation of animal populations, demonstrating its role in upper trophic marine species. We show that environmental variation leaves a signal in long-term population dynamics of seabirds with potentially important consequences for their demography and trophic interactions.

Species breeding at high latitudes face a significant challenge of surviving the winter. Such conditions are particularly severe for diurnal marine endotherms such as seabirds. A critical question is therefore what behavioural strategies such species adopt to maximise survival probability. We tested 3 hypotheses: (1) they migrate to lower latitudes to exploit longer day length (‘sun-chasing’), (2) they forage at night (‘night-feeding’), or (3) they target high-quality food patches to minimise foraging time (‘feasting’). We studied the winter migration and foraging strategies of European shags Phalacrocorax aristotelis from 6 colonies across a latitudinal gradient from temperate regions to north of the Arctic Circle using geolocators deployed over 11 winters. We found evidence for ‘sun-chasing’, whereby average southerly movements were greatest from colonies at higher latitudes. However, a proportion of individuals from higher latitudes remained resident in winter and, in the absence of daylight, they foraged during twilight and only very occasionally during the night. At lower latitudes, there was little evidence that individuals migrated south, nocturnal feeding was absent, and twilight feeding was infrequent, suggesting that there was sufficient daylight in winter. There was no evidence that winter foraging time was lowest at higher latitudes, as predicted by the ‘feasting’ hypothesis. Our results suggest that shags adopt different behavioural strategies to survive the winter across their latitudinal range, dictated by the differing light constraints. Our study highlights the value of multi-colony studies in testing key hypotheses to explain population persistence in seabird species that occur over large latitudinal ranges.

Seabirds are undergoing drastic declines globally and spend the non-breeding season at sea, making it challenging to study the drivers of their survival. Harsh weather and changes in climate conditions can have large impacts on seabird population dynamics through increased mortality. The intensity and persistence of extreme events are forecasted to increase with global warming. As shared conditions can induce population synchrony, multi-population studies of key demographic parameters are imperative to explore the influence of climate change. We used long-term mark-recapture data and position data to determine non-breeding stop-over areas of 5 Atlantic puffin (Fratercula arctica) populations over a latitudinal gradient in the Northeast Atlantic (56°11’-70°23’N). We investigated synchrony in adult survival in relation to shared stop-over areas. We quantified effects of extreme extra-tropical cyclones (ETCs) specific to populations’ stop-over areas and the North Atlantic Oscillation on adult survival. Populations with overlapping stop-over areas exhibited temporal synchrony in survival rates. Winter ETCs negatively influenced survival in 1 population, which was the one most exposed to extreme weather, but did not directly influence adult survival in the other 4 populations. Synchrony among populations with shared stop-over areas highlights the importance of these areas for adult survival, a key life-history rate. However, extreme weather was not identified as a driving factor for the majority of study populations. This suggests other factors in these areas, likely related to bottom-up trophic interactions, as environmental drivers of synchrony in the survival of Atlantic puffins.

Carry-over effects, whereby events in one season have consequences in subsequent seasons, have important demographic implications. Although most studies examine carry-over effects across 2 seasons in single populations, the effects may persist beyond the following season and vary across a species’ range. To assess potential carry-over effects across the annual cycle and among populations, we deployed geolocation loggers on black-legged kittiwakes Rissa tridactyla at 10 colonies in the north-east Atlantic and examined relationships between the timing and destination of migratory movements and breeding success in the year of deployment and sub - sequent season. Both successful and unsuccessful breeders wintered primarily in the north-west Atlantic. Breeding success affected the timing of migration, whereby unsuccessful breeders departed the colony earlier, arrived at the post-breeding and main wintering areas sooner, and departed later the following spring. However, these patterns were only apparent in colonies in the south-west of the study region. Furthermore, the effect of breeding success was stronger on migration timing in the first part of the winter than later. Timing of migratory movements was weakly linked to subsequent breeding success, and there was no detectable association between breeding success in the 2 seasons. Our results indicate temporal structure and spatial hetero - geneity in the strength of seasonal interactions among kittiwakes breeding in the north-east Atlantic. Variable fitness consequences for individuals from different colonies could have important implications for population processes across the species’ range and suggest that the spatiotemporal dynamics of carry-over effects warrant further study. Seasonal interactions · Migration · Reproduction · Life-history strategies · Geolocation · Black-legged kittiwake · Rissa tridactyla · North Atlantic

Aim: An understanding of the non-breeding distribution and ecology of migratory species is necessary for successful conservation. Many seabirds spend the non-breeding season far from land, and information on their distribution during this time is very limited. The black-legged kittiwake, Rissa tridactyla, is a widespread and numerous seabird in the North Atlantic and Pacific, but breeding populations throughout the Atlantic range have declined recently. To help understand the reasons for the declines, we tracked adults from colonies throughout the Atlantic range over the non-breeding season using light-based geolocation. Location: North Atlantic. Methods: Geolocation data loggers were deployed on breeding kittiwakes from 19 colonies in 2008 and 2009 and retrieved in 2009 and 2010. Data from 236 loggers were processed and plotted using GIS. Size and composition of wintering populations were estimated using information on breeding population size. Results: Most tracked birds spent the winter in the West Atlantic, between Newfoundland and the Mid-Atlantic Ridge, including in offshore, deep-water areas. Some birds (mainly local breeders) wintered in the North Sea and west of the British Isles. There was a large overlap in winter distributions of birds from different colonies, and colonies closer to each other showed larger overlap. We estimated that 80% of the 4.5 million adult kittiwakes in the Atlantic wintered west of the Mid-Atlantic Ridge, with only birds from Ireland and western Britain staying mainly on the European side. Main conclusions: The high degree of mixing in winter of kittiwakes breeding in various parts of the Atlantic range implies that the overall population could be sensitive to potentially deteriorating environmental conditions in the West Atlantic, e.g. owing to lack of food or pollution. Our approach to estimating the size and composition of wintering populations should contribute to improved management of birds faced with such challenges.

1. Determining the precise timing and location of major demographic bottlenecks, such as periods of low survival, is key to identifying ecological causes of variation in population growth rate. Such understanding is key to designing efficient and effective mitigation. 2. In a protected population of red-billed chough Pyrrhocorax pyrrhocorax on Islay, Scotland, variation in population growth rate largely reflects among-year variation in first-year survival. First-year survival was unprecedentedly low during 2007–2010, threatening population viability. 3. We used colour-ring resightings to estimate monthly survival probabilities (Φm) throughout the first year from fledging for eight chough cohorts (totalling 519 individuals) representing the full observed range of variation in first-year survival. We thereby identify the time and location of recent low survival. 4. On average across all cohorts, Φm varied among months, being low during the first month after ringing (May–June, accounting for c. 24% of all first-year mortality) and high during the last 4 months of the first year (January–May, accounting for c. 6% of all first-year mortality). Most mortality (c. 70%) occurred after fledglings dispersed from natal territories. 5. The 2007–2009 cohorts experienced low Φm during July–December. This represents an additional low survival period compared to previous cohorts rather than decreased Φm across all months or further decreases through periods when Φm was low across all cohorts. 6. Synthesis and applications. These data have general relevance in showing that dramatically low annual survival, which needs to spark rapid management action, can reflect different and unanticipated periods of low survival rather than exaggeration of typical variation. With specific regard to conserving Islay’s chough population, our data show that sub-adult survival has recently been low during July–December, probably reflecting conditions on key grassland foraging areas. Managers aiming to increase population viability should increase invertebrate diversity, abundance and availability at these times and locations, thereby increasing foraging options available to choughs.

An organism’s pattern of development can have important long-term fitness effects. In species where the sexes differ in size or other phenotypic traits, they may also have different optimal developmental rates. This influences both parental sex allocation strategies and susceptibility of the sexes to early developmental conditions. However, sex differences in developmental rate and vulnerability to environment during the embryonic period are not well understood. In birds, sibling competition and hatching asynchrony may select for accelerated embryonic development of the last offspring in order to reduce their competitive disadvantage after hatching. They may advance their hatching in response to vocal stimuli by the older siblings. It is, however, unclear whether this flexibility in developmental rates is sex specific. In this study, we experimentally manipulated between-embryo contact and tested whether this affected the pre-natal developmental rate and post-hatching performance of male and female offspring from last-laid eggs in the herring gull. Post-hatching performance was measured both in competitive and non-competitive situations. Among young incubated in isolation, males hatched faster than females, but both sexes fledged in similar, relatively good condition. Among young incubated with normal between-embryo contact, hatching time did not differ between sexes, but males fledged in poorer condition than females, regardless of whether they were reared singly or in a brood. These results suggest that male and female offspring differ in their ability to mitigate the costs of hatching asynchrony.

The quality of conditions provided by avian parents will have consequences for both parental and offspring fitness. While many components of avian reproduction appear to vary with parental age, the effect of age on incubation has largely been ignored so far. In this study, we tested whether young herring gulls provide a different incubation environment from mature ones and whether this has consequences for offspring performance. Laying and rearing conditions were standardised using a cross-fostering protocol. Egg predation rates tended to be higher in the nests of young parents. However, nest site, nest construction and egg temperature during incubation did not vary with parental age. Overall, the duration of incubation was shorter in young compared to mature birds and this reflected the later laying date of the former, since incubation duration generally decreased across the season. However, male eggs incubated by young parents had longer incubation periods than predicted for their laying dates. In contrast, incubation length of female eggs incubated by young pairs, and of male and female eggs incubated by mature birds did not deviate from the expected for any given laying date. Offspring that had been incubated by young parents had considerably poorer survival than those incubated by mature pairs, despite being reared under standardized, favourable conditions (singly, by mature parents). This was due to increased mortality among female chicks that had been incubated by young parents. The chicks incubated as eggs by young and mature birds, which survived until fledging, did not differ in body mass and size growth, or body condition. The results of this study demonstrate that parental age can influence offspring performance via variation in incubation environment, and that females are more susceptible than males to conditions experienced during embryonic development.

Summary 1Poorer performance in young compared with older age classes is common among iteroparous animals, and may result from age effects acting on different phases of reproduction. However, such effects have generally not been teased apart, and as a result their separate consequences for offspring fitness have remained unclear. 2We tested whether young and mature Herring Gulls produce offspring of different quality due to differences in their egg-production capacity. An experimental protocol was used to standardize incubation and rearing conditions. 3Eggs laid by young females were smaller than eggs of mature birds. They had a reduced absolute amount of albumen, which reflected mainly lower water content. 4Within the young age class, female eggs were smaller than male eggs and had significantly paler yolk. This was due to young birds having reduced water content of the albumen, as well as carotenoid content in eggs containing a female embryo. In the mature class there were no such differences in resource allocation in relation to embryo sex. The sex ratio of clutches did not vary with parental age. 5Hatchlings from eggs produced by young pairs were smaller skeletally but had similar mass to hatchlings from eggs laid by mature birds. When reared by mature foster parents, they did not differ in structural growth or survival. However, they showed a different pattern of growth in body mass. 6Our results suggest that poorer reproductive performance of young breeders is not simply a consequence of poorer egg quality, as this was largely compensated for when the eggs of young pairs were incubated and reared by mature foster parents. However, it is also possible that adverse effects of lower egg quality are not manifest in the offspring until later in life.

Marine spatial planning aims to deliver sustainable use of marine resources by minimizing environmental impacts of human activities and designating Marine Protected Areas. This poses a challenge where species’ distributions show spatio-temporal heterogeneity. However, due to logistic constraints and challenging timescales many studies of distribution are undertaken over few years or on a restricted subset of the population. Long-term studies can help identify the degree of uncertainty in those less comprehensive in space and time. We quantify inter-annual and sub-colony variation in the summer foraging distribution of a population of European shags Phalacrocorax aristotelis, using a tracking data set comprising 320 individuals and 1106 foraging trips in 15 years from 1987 to 2010. Foraging distribution over the study period was concentrated in three areas. Data from one and two years captured an average of 54% and 64% of this distribution, respectively, but it required 8 years’ data to capture over 90% of the distribution. Foraging range increased with population size when breeding success was low, suggesting interplay between extrinsic and intrinsic effects. Furthermore, females had foraging ranges on average 36% greater than males. Finally, sub-colony segregation occurred in foraging areas up to 4 km from the colony and in the most distant locations (>10 km), whilst there was considerable overlap at intermediate distances (6–10 km). Our study highlights important considerations for marine spatial planning in particular, and species conservation in general, notably the proportion of the population distribution identified, the prevailing conditions experienced and the need for balanced sampling across the population. 2014 The Authors. Published by Elsevier Ltd. This is an open access article under the CC BY license

Miniature geolocator loggers (Global Location Sensing, GLS) that provide daily locations of birds have revolutionised the study of winter ecology and migration patterns of seabirds. A long-term study of ringing recoveries and analyses of heavy metals and pollutants in tissues of Atlantic puffins Fratercula arctica from the Isle of May, south-east Scotland, suggested that this population wintered mainly within the North Sea. However, deployment of GLS devices over the 2007/2008 winter showed that many breeding birds made major excursions into the east Atlantic. This winter was the second of two when survival was extremely low (survival in 2006/2007 and 2007/2008 was 0.696 and 0.695, respectively, compared to the average of 0.922 over the period 1984/1985–2005/2006). These low rates of survival suggested that the unexpected use of the Atlantic might have been associated with unusually poor conditions in the North Sea as indicated by very low breeding success in 2007. Survival rate returned to previous levels in 2008/2009 providing the opportunity to test whether higher survival was associated with birds remaining in the North Sea, or whether movements into the Atlantic are a feature of this population unrelated to survival. Accordingly, geolocators were deployed over the 2009/2010 winter when adult survival was subsequently established to be high (0.913). We found greater support for the hypothesis that winter distribution is not associated with survival. Thus, 8 (40 %) of 20 individuals followed in 2009/2010 went into the Atlantic, a rate not significantly different from 11 (58 %) of the 19 followed in the 2007/2008 winter. Indeed, birds actually spent longer in the Atlantic and used a wider variety of areas in 2009/2010, although the time spent away from the colony was significantly shorter than in 2007/2008. Since our data were from individuals that survived, remaining in or moving out of the North Sea can both be successful strategies during winters when the population as a whole shows either high or low survival rates. Unfortunately, we do not know where birds that died had gone, and hence, the relative survival of birds that did or did not move into the Atlantic. Determining the link between survival and wintering area for any seabird remains a formidable challenge and will have to await the development of technologies that can determine both where and when birds die.

Relationships between events in one period of the annual cycle and behaviour in subsequent seasons are key determinants of individual life histories and population dynamics. However, studying such associations is challenging, given the difficulties in following individuals across seasons, particularly in migratory species. Relationships between breeding performance and subsequent winter ecology are particularly poorly understood, yet are likely to be profoundly important because of the costs of reproduction. Using geolocation technology, we show that black-legged kittiwakes that experienced breeding failure left their colony in southeast Scotland earlier than successful breeders. Moreover, a greater proportion of unsuccessful breeders (94% versus 53% successful) travelled over 3000 km to the West Atlantic, whereas fewer visited the East Atlantic (31% versus 80% successful), less than 1000 km from the colony. The two groups did not differ in the timing of return to the colony the following spring. However, 58 per cent of males made a previously undescribed long-distance pre-breeding movement to the central Atlantic. Our results demonstrate important links between reproductive performance and winter distribution, with significant implications for population dynamics. Furthermore, macro-scale segregation associated with breeding outcome is relevant to defining important wintering areas, in particular among declining species experiencing increasingly regular breeding failure.

1. Understanding the degree to which reproductive success varies with an individual's age and lifespan, and the degree to which population-level variation mirrors individual-level variation, is central to understanding life-history evolution and the dynamics of age-structured populations. We quantified variation in the survival probability of offspring, one key component of reproductive success and fitness, in relation to parent age and lifespan in a wild population of red-billed choughs (Pyrrhocorax pyrrhocorax). 2. On average across the study population, the first-year survival probability of offspring decreased with increasing parent age and lifespan; offspring of old parents were less likely to survive than offspring of young parents, and offspring of long-lived parents were less likely to survive than offspring of short-lived parents. 3. However, survival did not vary with parent age across offspring produced by groups of parents that ultimately had similar lifespans. 4. Rather, across offspring produced by young parents, offspring survival decreased with increasing parent lifespan; parents that ultimately had long lifespans produced offspring that survived poorly, even when these parents were breeding at young ages. 5. The average decrease in offspring survival with increasing parent age observed across the population therefore reflected the gradual disappearance of short-lived parents that produced offspring that survived well, not age-specific variation in offspring survival within individual parents. 6. The negative correlation between offspring survival and maternal lifespan was strongest when environmental conditions meant that offspring survival was low across the population. 7. These data suggest an environment-dependent trade-off between parent and offspring survival, show consistent individual variation in the resolution of this trade-off that is set early in a parent's life, and demonstrate that such structured life-history variation can generate spurious evidence of senescence in key fitness components when measured across a population.

1. Quantifying the pattern of temporal and spatial variation in demography, and identifying the factors that cause this variation, are essential steps towards understanding the structure and dynamics of any population. 2. One critical but understudied demographic rate is pre-breeding survival. We used long-term colour-ringing data to quantify temporal (among-year) and spatial (among-nest site) variation in pre-breeding survival in red-billed choughs (Pyrrhocorax pyrrhocorax) inhabiting Islay, Scotland, and identified environmental correlates of this variation. 3. Random-effects capture-mark-recapture models demonstrated substantial temporal and spatial process variance in first-year survival; survival from fledging to age 1 year varied markedly among choughs fledged in different years and fledged from different nest sites. Spatial variance exceeded temporal variance across choughs fledged from well-studied nest sites. 4. The best-supported models of temporal variation suggested that first-year survival was higher in years following high tipulid larvae abundance and when weather conditions favoured increased invertebrate productivity and/or availability to foraging choughs. These variables explained up to 80% of estimated temporal process variance. 5. The best-supported models of spatial variation suggested that first-year survival was higher in choughs fledged from nest sites that were further from exposed coasts and closer to flocking areas, and surrounded by better habitat and higher chough density. These variables explained up to 40% of estimated spatial process variance. 6. Importantly, spatio-temporal models indicated interactive effects of weather, tipulid abundance, local habitat and local chough density on first-year survival, suggesting that detrimental effects of poor weather and low tipulid abundance may be reduced in choughs fledged from nest sites surrounded by better foraging habitat and lower chough density. 7. These analyses demonstrate substantial temporal and small-scale spatial variation in pre-breeding survival, a key demographic rate, and indicate that this variation may reflect interactive effects of weather, prey abundance, habitat and geography. These patterns illustrate the value of holistic models of demographic variation, and indicate environmental factors that may limit the growth rate of Islay's protected chough population.