Ecosystem structure and biophysical processes mediate biological responses to climate changes, but few studies have examined impacts of this dynamic among upper trophic levels. We investigated ecosystem differences in how diverse seabird populations across the northern hemisphere have responded to changes in regional mixed layer temperature and water column stratification. Using 138 time series of breeding productivity over the past half-century, we show that seabird reproductive productivity has declined in the Arctic and North Atlantic but not in the Pacific during a period of ubiquitous mixed layer warming and regionally-variable stratification trends. Models of breeding productivity and ocean drivers show that seabird responses to climate change vary by ecosystem. Additionally, ecosystems in which seabirds exhibit detectibly declining productivity tend to have lower overall diet diversity across seabird species. These findings emphasize the importance of ecosystem processes and structure in determining the vulnerability of marine predators to climate change.

AimEcologically similar species living in sympatry are expected to segregate to reduce the effects of competition where resources are limiting. Segregation from heterospecifics commonly occurs in space, but it is often unknown whether such segregation has underlying environmental causes. Indeed, species could segregate because of different fundamental environmental requirements (i.e., 'niche divergence'), because competitive exclusion at sympatric sites can force species to either change the habitat use they would have at allopatric sites (i.e., 'niche displacement') or to avoid certain areas, independently of habitat (i.e., 'spatial avoidance'). Testing these hypotheses requires the comparison between sympatric and allopatric sites. Understanding the competitive mechanisms that underlie patterns of spatial segregation could improve predictions of species responses to environmental change, as competition might exacerbate the effects of environmental change.LocationNorth Atlantic and Arctic.TaxaCommon guillemots Uria aalge and Br & uuml;nnich's guillemots Uria lomvia.MethodsHere, we examine support for these explanations for spatial segregation in two closely-related seabird species, common guillemots (Uria aalge) and Br & uuml;nnich's guillemots (U. lomvia). For this, we collated a pan-Atlantic data set of breeding season foraging tracks from 1046 individuals, collected from 20 colonies (8 sympatric and 12 allopatric). These were analysed with habitat models in a spatially transferable framework to compare habitat preferences between species at sympatric and allopatric sites.ResultsWe found no effect of the distribution of heterospecifics on local habitat preferences of the focal species. We found differences in habitat preferences between species, but these were not sufficient to explain the observed levels of spatial segregation at sympatric sites.Main ConclusionsAssuming we did not omit any relevant environmental variables, these results suggest a mix of niche divergence and spatial avoidance produces the observed patterns of spatial segregation.

Surveys of seabirds breeding along the coast of Sør-Varanger, NE Norway were carried out in 1989, 1996, 1999 and 2002 as a continuation of a monitoring study started in 1970. Until 1983, by which time 47 colonies were occupied, there had been a rapid increase in numbers of Black-legged Kittiwakes Rissa tridactyla and European Shags Phalacrocorax aristotelis breeding in the area while the Great Cormorant P. carbo population had first increased and then declined. Between 1983 and 2002, the Kittiwake and Shag populations declined by ca. 60% and 85% respectively while that of Cormorants first continued to decline and then recovered to levels recorded in the 1970s. The decline in Common Guillemot Uria aalge and Razorbill Alca torda numbers recorded since 1970 culminated in 1989 when no birds were seen in the region. Since then, one individual Common Guillemot was seen in 1999. In 2002 approximately 7500 pairs of Kittiwake, 770 pairs of Cormorant and 15 pairs of Shag bred in 23 colonies. Possible causes for the main population changes are briefly discussed.

The breeding populations of Atlantic Puffins Fratercula arctica, Common Guillemots Uria aalge and Black-legged Kittiwakes Rissa tridactyla have been monitored on Hornøya, East Finnmark since 1980 as part of the Norwegian seabird monitoring programme. Whereas numbers of Puffin burrows in plots monitored in 1981 - 1993 increased at a rate of 2.6% per year, there was no subsequent trend in counts made in a new scheme of circular plots started in 1990. Numbers of Common Guillemots collapsed between 1985 - 1987 but have since increased in all plots at a rate of 11.6% per year. Kittiwake numbers dropped significantly (1.5% per year) between 1980 - 1994, but stabilised between 1994 - 2000. Suggestions are made for improvements to the monitoring scheme to overcome inconsistencies in changes in numbers between plots (e.g. of Kittiwakes), to increase the sensitivity of the counts and for additions to help explain why any changes in the populations occur.

Forecasting population responses to rapidly changing marine ecosystems requires mechanistic models integrating complex demographic processes, fitted to long time series, across large spatial scales. We used a Bayesian metapopulation model fit to colony census data and climatic covariates spanning 1900-2100 for all Northeast Atlantic colonies of an exemplar seabird, the Northern gannet (Morus bassanus) to investigate metapopulation dynamics under two climate scenarios. Fecundity varied non-linearly with near-surface air temperature and recruitment was depressed by sea surface temperature. We predict regime changes in density dependence as marine carrying capacities become constrained with increasing SST. Sensitivity to climate change varied across space and time, disadvantaging southwestern colonies whilst benefitting northern ones. Such sensitivity is noteworthy for a species previously assumed robust to climate change. We provide a spatial overview of climate sensitivities across a metapopulation to help with evidence-based conservation management and open the way for similar mechanistic explorations for other colonial species.

Timing of moult of the inner primaries by breeding Black-legged Kittiwakes is very similar among colonies ranging from southern England to northern Norway, but can be much later at very high-latitude colonies, such as at Svalbard, where breeding is delayed by the late spring thaw. Breeding adult Black-legged Kittiwakes almost all replaced the first and second (innermost) primary feathers on their wings while breeding.

change affects timing of reproduction in many

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 change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

Density-dependent feedback is recognized as important regulatory mechanisms of population size. Considering the spatial scales over which such feedback operates has advanced our theoretical understanding of metapopulation dynamics. Yet, metapopulation models are rarely fit to time-series data and tend to omit details of the natural history and behavior of long-lived, highly mobile species such as colonial mammals and birds. Seabird metapopulations consist of breeding colonies that are connected across large spatial scales, within a heterogeneous marine environment that is increasingly affected by anthropogenic disturbance. Currently, we know little about the strength and spatial scale of density-dependent regulation and connectivity between colonies. Thus, many important seabird conservation and management decisions rely on outdated assumptions of closed populations that lack density-dependent regulation. We investigated metapopulation dynamics and connectivity in an exemplar seabird species, the Northern gannet (Morus bassanus), using more than a century of census data of breeding colonies distributed across the Northeast Atlantic. We developed and fitted these data to a novel hierarchical Bayesian state-space model, to compare increasingly complex scenarios of metapopulation regulation through lagged, local, regional, and global density dependence, as well as different mechanisms for immigration. Models with conspecific attraction fit the data better than the equipartitioning of immigrants. Considering local and regional density dependence jointly improved model fit slightly, but importantly, future colony size projections based on different mechanistic regulatory scenarios varied widely: a model with local and regional dynamics estimated a lower metapopulation capacity (645,655 Apparently Occupied Site [AOS]) and consequently higher present saturation (63%) than a model with local density dependence (1,367,352 AOS, 34%). Our findings suggest that metapopulation regulation in the gannet is more complex than traditionally assumed, and highlight the importance of using models that consider colony connectivity and regional dynamics for conservation management applications guided by precautionary principles. Our study advances our understanding of metapopulation dynamics in long-lived colonial species and our approach provides a template for the development of metapopulation models for colonially living birds and mammals.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

Two subspecies of the Great Cormorant breed in Norway, the continental Phalacrocorax carbo sinensis in the south, along the Skagerrak coast, and the marine P. c. carbo from central Norway and northwards. Here we review the information existing until 2017 on population status and trends, breeding performance and diet of these two subspecies in Norway. The most recent national population estimates are approximately 2500 (in 2012) and 19,000 (in 2012–2014) breeding pairs of sinensis and carbo, respectively. The sinensis population established itself in 1996 in Rogaland at the south-western tip of Norway, and in 1997 in Østfold close to the Swedish border; in both areas it increased for about ten years. Since then, the numbers have stabilised. For carbo, the population increased from 21,000 pairs in the early 1980s to 27,000 in 1995, and then decreased to the current number of 19,000 pairs. Significant annual variations in clutch size and reproductive output have been observed, but the drivers of these changes have not been identified. Unidentified gadoids and Atlantic Cod Gadus morhua were the most common prey of carbo, whereas inshore species such as Corkwing Wrasse Symphodus melops, Rockcook Centrolabrus exoletus, Goldsinny Wrasse Ctenolabrus rupestris and Black Goby Gobius niger were the most common prey in the eastern Skagerrak caught by sinensis. Carbo took very large numbers of 1–3-year-old gadoids during the year, and we cannot exclude the possibility this can have local effects on fish mortality rates.

The conservation of migratory marine species, including pelagic seabirds, is challenging because their movements span vast distances frequently beyond national jurisdictions. Here, we aim to identify important aggregations of seabirds in the North Atlantic to inform ongoing regional conservation efforts. Using tracking, phenology, and population data, we mapped the abundance and diversity of 21 seabird species. This revealed a major hotspot associated with a discrete area of the subpolar frontal zone, used annually by 2.9–5 million seabirds from ≥56 colonies in the Atlantic: the first time this magnitude of seabird concentrations has been documented in the high seas. The hotspot is temporally stable and amenable to site‐based conservation and is under consideration as a marine protected area by the OSPAR Commission. Protection could help mitigate current and future threats facing species in the area. Overall, our approach provides an exemplar data‐driven pathway for future conservation efforts on the high seas.

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.

Climate change and other human activities are causing profound effects on marine ecosystem productivity. We show that the breeding success of seabirds is tracking hemispheric differences in ocean warming and human impacts, with the strongest effects on fish-eating, surface-foraging species in the north. Hemispheric asymmetry suggests the need for ocean management at hemispheric scales. For the north, tactical, climate-based recovery plans for forage fish resources are needed to recover seabird breeding productivity. In the south, lower-magnitude change in seabird productivity presents opportunities for strategic management approaches such as large marine protected areas to sustain food webs and maintain predator productivity. Global monitoring of seabird productivity enables the detection of ecosystem change in remote regions and contributes to our understanding of marine climate impacts on ecosystems.

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.

The timing of annual events such as reproduction is a critical component of how free‐living organisms respond to ongoing climate change. This may be especially true in the Arctic, which is disproportionally impacted by climate warming. Here, we show that Arctic seabirds responded to climate change by moving the start of their reproduction earlier, coincident with an advancing onset of spring and that their response is phylogenetically and spatially structured. The phylogenetic signal is likely driven by seabird foraging behavior. Surface‐feeding species advanced their reproduction in the last 35 years while diving species showed remarkably stable breeding timing. The earlier reproduction for Arctic surface‐feeding birds was significant in the Pacific only, where spring advancement was most pronounced. In both the Atlantic and Pacific, seabirds with a long breeding season showed a greater response to the advancement of spring than seabirds with a short breeding season. Our results emphasize that spatial variation, phylogeny, and life history are important considerations in seabird phenological response to climate change and highlight the key role played by the species’ foraging behavior.

To identify priorities for management of seabirds during the breeding season, it is important to understand the ecological mechanisms driving chick growth and survival. In this study, we examined the effects of diet and prevailing weather on the growth and survival of chicks of black-legged kittiwakes Rissa tridactyla over a 10 yr period at Anda, a seabird colony in northern Norway. We show that across all years, there was a significant effect of diet composition delivered to chicks on their growth and survival. A higher proportion of sandeel Ammodytes spp. in the chick diet was associated with an increase in daily growth rates, a pattern that was especially pronounced for the youngest chick in 2-chick broods. A high proportion of mesopelagic fish in the chick diet was associated with a decrease in survival, again, especially for the youngest chick in 2-chick broods. Periods of strong southerly winds also led to reduced survival, probably linked to nests being washed down from the colony. Growth rates of kittiwake chicks were negatively affected by wind speed, likely due to adults having to work more in the exposed habitats in strong winds, causing a reduction in the amount of food supplied to the chicks. Our results emphasise the importance of conservation of specific marine habitats shown to be important foraging areas in ensuring the reproductive success of seabirds. This might prove increasingly important if future climate regimes make ecological conditions more challenging for seabirds.

Reproductive timing in many taxa plays a key role in determining breeding productivity 1 , and is often sensitive to climatic conditions 2 . Current climate change may alter the timing of breeding at different rates across trophic levels, potentially resulting in temporal mismatch between the resource requirements of predators and their prey 3 . This is of particular concern for higher-trophic-level organisms, whose longer generation times confer a lower rate of evolutionary rescue than primary producers or consumers 4 . However, the disconnection between studies of ecological change in marine systems makes it difficult to detect general changes in the timing of reproduction 5 . Here, we use a comprehensive meta-analysis of 209 phenological time series from 145 breeding populations to show that, on average, seabird populations worldwide have not adjusted their breeding seasons over time (−0.020 days yr−1) or in response to sea surface temperature (SST) (−0.272 days °C−1) between 1952 and 2015. However, marked between-year variation in timing observed in resident species and some Pelecaniformes and Suliformes (cormorants, gannets and boobies) may imply that timing, in some cases, is affected by unmeasured environmental conditions. This limited temperature-mediated plasticity of reproductive timing in seabirds potentially makes these top predators highly vulnerable to future mismatch with lower-trophic-level resources 2 . Time of reproduction may be altered as the climate changes. For seabirds, it is shown that there has not been an adjustment in timing as the climate changes and the sea surface warms. This lack of plasticity could result in a mismatch with food resources.

Birds allocate substantial resources to their eggs during the laying period, resources also needed for other concurrent costly processes such as mate acquisition, nest building and site defence. Egg and clutch sizes may thus vary in response to food availability prior to egg laying. We investigated the variation in egg and clutch size of black-legged kittiwakes Rissa tridactyla in a North Norwegian colony over a 33 yr period (1980-2012). Considerable interannual variation was evident in both egg and clutch sizes, but no temporal trends were observed. To identify environmental conditions explaining the variation, we modelled egg size and number in relation to the influx of warm Atlantic Water (AW) into the Barents Sea and to the abundance of 2 of the kittiwakes' most preferred prey species. Most of the variation was explained by the volumes of AW that flowed into the Barents Sea in winter and autumn. Both had a negative effect on egg investment. There was also a smaller, positive effect on egg investment of AW inflow (and capelin numbers for clutch size) immediately prior to egg laying. The negative impact of an increased influx of AW on kittiwakes may be partly due to the resulting warming of the waters, forcing a more northerly distribution of capelin beyond the foraging range of the kittiwakes at the colony or to changes at different trophic levels that are detrimental to kittiwake forage fish ecology. A further warming of the Barents Sea through climate change is thus forecast to be detrimental for kittiwakes.