Colonial Seabirds Research Articles

Competition and Facilitation Influence Central Place Foraging Ecology in a Colonial Marine Predator.

Coloniality is strongly shaped by aspects of social foraging behaviour. For example, colonies may be important sources of information, while food competition may increase foraging efforts and limit colony size. Understanding foraging ecology considering these apparent trade-offs is required to develop a better understanding of colonial living. We combined animal-borne GPS, cameras and dive recorders to study social foraging in breeding adult northern gannets Morus bassanus-a wide-ranging colonial seabird. We first tested for indirect evidence of prey depletion around the colony by estimating dive location, depth and duration. Next, we tested for sociality during different behaviours (commuting, foraging and resting) and distance from the colony. Finally, we quantified flocks of inbound and outbound birds to compare social foraging between outbound and inbound legs of the commute. Dive probability and depth (n = 46 individuals; n = 1590 dives) increased with distance from the colony, creating dive clusters at ~100 and 180 km consistent with conspecific prey depletion. Camera stills (n = 8 individuals; n = 7495 images) show gannets are highly social, but this varied among behaviours. Sociality was highest during foraging and commuting; especially inbound and social foraging was more likely far from the colony. Gannets were equally likely to be solitary or social when leaving the colony but returning birds were more likely in larger flocks. In summary, despite experiencing intraspecific competition for food, gannets engage in dynamic, context-dependent social foraging associations. Conspecifics aggregated far from the colony possibly because of a prey depletion halo closer to home, but this provided potential benefits via local enhancement and by returning to the colony in flocks. Our results therefore illustrate how competition may, paradoxically, facilitate some aspects of group foraging in colonial animals.

Population monitoring and conservation implications of intra- and interspecific nest occupation rates in swallows.

With the exception of a few groups of birds, such as large raptors and colonial seabirds, direct counts of nests cannot be conducted over very large areas for most of the abundant and widely distributed species, and thus indirect methods are used to estimate their relative abundances and population sizes. However, many species of the Family Hirundinidae (swallows and martins) build their mud nests in discrete, predictable and accessible sites, which are reused across years. Therefore, the direct count of active nests could constitute a reliable method for estimating breeding population sizes and their changes at large spatial and temporal scales. We illustrate the feasibility of this monitoring approach through a single year survey of >2700 nests of three coexisting Old-World species, the barn swallow (Hirundo rustica), the red-rumped swallow (Cecropis daurica), and the crag martin (Ptyonoprogne rupestris), distributed across Portugal and Spain. Our results revealed changes in the use of nesting substrates and increases in interspecific nest usurpation rates over recent decades. While 56% of the nests of C. daurica were located in rocks five decades ago, almost 100% are nowadays located in anthropogenic substrates such as bridges, road culverts, and abandoned buildings, which could have favored the range expansion of this species. Nest occupation rates were surprisingly low (12% in C. daurica, 21% in H. rustica, and 37% in P. rupestris), and the proportion of abandoned nesting sites was very high (65% in C. daurica, 50% in H. rustica, and 27% in P. rupestris). Abandonment rates reflect the population decline reported for H. rustica. Notably, the usurpation of nests of C. daurica by house sparrows Passer domesticus, which is the main cause of breeding failure, has increased from 2.4% in 1976-1979 to 34.7% of the nests nowadays. The long-term monitoring of nests may constitute a reliable and affordable method, with the help of citizen science, for assessing changes in breeding population sizes and conservation threats of these and other mud-nest building hirundines worldwide.

Social, not genetic, programming of development and stress physiology of a colonial seabird.

Phenotypic differences often stem from genetic/maternal differences and/or early-life adaptations to local environmental conditions. In colonial animals, little is known on how variation in the social environment is embedded into individual phenotypes, nor what the consequences are on individual fitness. We conducted an experimental cross-fostering study on king penguins (Aptenodytes patagonicus), exchanging eggs among 134 pairs breeding in high-density (67 pairs) or low-density (67 pairs) areas of the same breeding colony. We investigated differences in parent and chick phenotypes and survival in relation to the density of their origin and foster environment. Adults breeding in colony areas of high density exhibited decreased resting behaviour and increased aggression and vigilance, increased hypometabolism during incubation fasts, and more moderate corticosterone responses shaped by exposure to chronic stressors (e.g. constant aggression by neighbours). Chick phenotypes were more influenced by the environment in which they were raised than their genetic/maternal origin. Chicks raised in high-density colonial environments showed enhanced weight gain and survival rates regardless of the density of their genetic parents' breeding areas. Our study experimentally shows advantages to breeding in colonial areas of higher breeder densities in king penguins, and highlights the importance of social settings in shaping phenotype expression in colonial seabirds.

Mercury bioaccumulation in three colonial seabird species in the Gulf of Maine

The methylated form of mercury, MeHg, is a neurotoxin that bioaccumulates and biomagnifies through aquatic food webs, reaching high concentrations in top trophic species. Many seabird species are wide-ranging and feed on forage fish, so they can be used as sentinel species to assess the level of mercury in pelagic or coastal food webs because they integrate the signal from large areas and from lower trophic levels. The Gulf of Maine provides habitat for many seabirds, including endangered roseate terns (Sterna dougalii), common terns (Sterna hirundo), and the southernmost breeding population of black guillemots (Cepphus grylle). Hg levels were assessed in down of newly hatched chicks of three seabird species to determine pre-hatching Hg exposure. Stable isotopes (δ15N, δ13C) in down and chick contour feathers grown after hatching were used as indicators of adult female diet in the period before laying the egg (down) and pre-fledging chick diet (contour feathers). Black guillemot down THg concentrations were 10.07 ± 2.88 μg/g (mean ± 1SD), 5.5× higher than common tern down (1.82 ± 0.436 μg /g), and 7.4× higher than roseate tern down (1.37 ± 0.518 μg/g). Black guillemots also had higher down feather δ15N values (15.1 ± 0.52 ‰) compared to common (13.0 ± 0.72 ‰) or roseate terns (12.8 ± 0.25 ‰), and in black guillemot down feathers, higher Hg concentrations were correlated with δ15N, an indicator of trophic level. Repeated testing of the same tissue types across multiple years is needed to monitor THg exposure for seabirds in the Gulf of Maine; additionally, monitoring species composition and Hg presence in prey species of the black guillemot population would help to determine the source of high THg concentrations in this species.

Using soundscape to monitor population size, demographics, and antipredator behavior in a dense aggregation of colonial seabirds

Migratory seabirds are vulnerable to decline due to climate change and anthropogenic disturbances. Common terns (Sterna hirundo) are highly vocal colonial seabirds that serve as bioindicators of their foraging grounds throughout their migratory range. Historically, monitoring colonial seabirds is invasive and time-consuming, and traditional acoustic approaches are complicated by high amounts of call overlap. Monitoring the behavioral ramifications of disturbance, as well as overall colony size and health, is crucial to implementing effective management decisions. However, methods are needed to do so efficiently and with minimal disturbance. In this study, we demonstrate that population size, demographics, and behavior can be assessed acoustically through changes in acoustic energy across varying temporal scales. To do this, we compared acoustic energy to in-person observations of nest density, chick-hatching, and investigator disturbance. We found that trends in acoustic energy align with observations of nest density, and the distribution of acoustic energy across frequency bands is indicative of colony demographics. Furthermore, we found a significant relationship between acoustic energy and investigator disturbance within 20 meters of an acoustic recorder. Overall, our findings suggest that colony-wide trends in population size, demographics, and behavior can be monitored via acoustic energy without the time-consuming analysis of individual calls.

Calls of the little auk (Alle alle) chicks reflect their behavioural contexts.

Animal vocalisations can often inform conspecifics about the behavioural context of production and the underlying affective states, hence revealing whether a situation should be approached or avoided. While this is particularly important for socially complex species, little is known about affective expression in wild colonial animals, and even less to about their young. We studied vocalisations of the little auk (Alle alle) chicks in the Hornsund breeding colony, Svalbard. Little auks are highly colonial seabirds, and adults convey complex behavioural contexts through their calls. We recorded chick calls during two contexts of opposite affective valence: handing by a human, and while they interact with their parents inside the nest. Using permuted discriminant function analysis and a series of linear mixed models, we examined the effect of the production context/associated affective valence on the acoustic parameters of those calls. Calls were reliably classified to their context, with over 97% accuracy. Calls uttered during handling had higher mean entropy, fundamental frequency, as well as lower spectral centre of gravity and a less steep spectral slope compared to calls produced during interactions with a parent inside the nest. The individuality of handling calls, assessed by information content, was lower than the individuality of calls uttered in the nest. These findings suggest that seabird chicks can effectively communicate behavioural/affective contexts through calls, conveying socially important messages early in development. Our results are mostly in line with emotional expression patterns observed across taxa, supporting their evolutionary continuity.

Highly pathogenic avian influenza causes mass mortality in Sandwich TernThalasseus sandvicensisbreeding colonies across north-western Europe

SummaryIn 2022, highly pathogenic avian influenza (HPAI) A(H5N1) virus clade 2.3.4.4b became enzootic and caused mass mortality in Sandwich TernThalasseus sandvicensisand other seabird species across north-western Europe. We present data on the characteristics of the spread of the virus between and within breeding colonies and the number of dead adult Sandwich Terns recorded at breeding sites throughout north-western Europe. Within two months of the first reported mortalities, 20,531 adult Sandwich Terns were found dead, which is >17% of the total north-western European breeding population. This is probably an under-representation of total mortality, as many carcasses are likely to have gone unnoticed and unreported. Within affected colonies, almost all chicks died. After the peak of the outbreak, in a colony established by late breeders, 25.7% of tested adults showed immunity to HPAI subtype H5. Removal of carcasses was associated with lower levels of mortality at affected colonies. More research on the sources and modes of transmission, incubation times, effective containment, and immunity is urgently needed to combat this major threat for colonial seabirds.

Movement patterns of foraging common terns (Sterna hirundo) breeding in an urban environment in coastal Virginia.

Nesting colonial seabirds are prime examples of central-place foragers, animals that must return to a central location (e.g., a breeding colony) after each bout of foraging. They must balance the costs and benefits of foraging with the need to return to their colonies frequently to form pair bonds during courtship, incubate, provision mates and offspring, and protect and rear young. For some populations, the loss and degradation of suitable breeding habitat due to human activities have necessitated the construction of new breeding sites and/or the restoration of previously occupied sites. South Island, which is part of the Hampton Roads Bridge-Tunnel (HRBT) complex in the Commonwealth of Virginia, U.S.A., is a human-created island that supported Virginia's largest mixed species seabird colony until 2020, when the expansion of the HRBT began and when all nesting seabirds were permanently excluded from the site. We studied the movement patterns of foraging common terns (Sterna hirundo) to determine how travel to and around foraging sites related to their colony location and to inform the siting and construction of a new breeding island. We tracked 18 individual common terns from 07 June to 29 June 2018, and we used a hidden Markov model to assign behavioral states and investigate common tern movements around the HRBT. Common terns spent more than half their time in the colony (58%), followed by time devoted to foraging (22%), and the remainder of their time was spent on outbound (15%) and inbound (5%) transit. Terns traveled as far as 98km from the colony, but on average foraged relatively close to South Island (13.6 ± 0.3km, mean ± 1 SD). Individuals tended to forage in the same locations, but there was variation among individuals. Flying to foraging sites uses energy during the already energetically costly breeding season, thus managers should prioritize placing a new colony site in a location that minimizes the distance traveled to the foraging locations frequented by the South Island birds while accounting for other life-history characteristics. These findings could help in the design and construction of new breeding sites or the restoration of current sites for other, related species, particularly for which these data do not exist.

Soils of the island bars of the Ola lagoon (Okhotsk sea)

The aim of the study. The purpose of the work is investigating soil formation on the island bars of the Ola Lagoon, as soil formation there has a special character due to the severe climate, dynamism and small age of the landscapes, and the influence of marine colonial birds. Location and time of the study. Field studies were conducted in June-July 2022 in the coastal zone of the Ola Lagoon of Magadan Region; laboratory and desk studies were conducted in 2022 and 2023. Methods. Methods included route observations with soil profile descriptions, mesomorphological studies; soil physical and chemical analyses: pH, electrical conductivity (TDS), loss on ignition (LOI); satellite photos interpretation and soil mapping. Results. The bar soils were qualified as dry-peat lithozems, differing in the thickness of the dry peat horizon TJ and the abundance of skeletal material (pebble). The dry-peat horizon is gradually merging into parent rock - dense marine pebbles, which are sometimes found in the dry-peat horizon due to storm "surges" from the sea. The TJ horizon consists of mesophilic plant residues and morphons of dark fine-disperse matter and a large number of semi-decomposed and living roots penetrating into the pebble layer. In the WRB system, soils correspond to Histic Leptosols (if shallow), or to Mawic Fibric Histosols. Conclusion. Formation of the dry-peat lithozems with the TJ-(TJ/С)-C profile is determined by the dry marine climate and low plant diversity of the meadows dominated by reed grass and wild rye. The main factors, shaping soil properties, are the island bars dynamics, specific of pebbles a parent rock material, salt impulverization and marine birds colonies presence. The research expands the knowledge on poorly studied soils of the region, and may contribute to substantiating their position in the classification of soils of Russia; in addition, the data obtained are useful within the framework of sustainable development, as it serves as nesting ground for colonial seabirds.

Mine or my neighbours’ offspring: an experimental study on parental discrimination of offspring in a colonial seabird, the little auk Allealle

Individual recognition (IR) abilities may result from various ecological and naturally selected features of a species. Complex IR mechanisms should develop when the risk of misidentification of a chick is high. For colonial seabirds, the ability to identify their own brood is crucial to ensure parental fitness. Vocalizations seem to be a key component of most parent–offspring interactions, although few studies have assessed the interindividual differences in seabird chick calls and their potential usage in IR. The little auk (Allealle), which breeds in dense colonies, constitutes a perfect model for testing IR. In this study, we (1) examined chick calls at different stages of the nesting period, and (2) cross-fostered chicks to examine the rate of acceptance/nonacceptance of chicks by parents. We found significant interindividual differences in chick begging and fledging calls. Surprisingly, all cross-fostered chicks in our experiments were accepted by their foster parents, and male parents were as equally likely to accept cross-fostered chicks as females, even though the sexes would be expected to differ in offspring recognition due to different postfledging interactions with the chick. The revealed individuality of chick calls suggests the potential for chick vocal recognition in the studied species, but parent birds may disregard the individual characteristics enabling chick discrimination. This may take place as long as the chick is found in the nest because of the high likelihood that the chick present there is the focal one. However, IR during and after fledging requires further study. Studying the complexity of IR mechanisms is important for better understanding various avian social relationships and interactions.

Disentangling the “many-eyes”, “dilution effect”, “selfish herd”, and “distracted prey” hypotheses in shaping alert and flight initiation distance in a colonial seabird

Group living is thought to have important antipredator benefits for animals, owing to the mechanisms of shared vigilance (“many-eyes” hypothesis), risk dilution (“dilution effect” hypothesis), and relative safety in the center of the group (“selfish herd” hypothesis). However, it can also incur costs since social stimuli, such as conspecific aggression, may distract individuals from anti-predator behavior (“distracted prey” hypothesis). We simultaneously evaluated how these four different hypotheses shape anti-predator behaviors of breeding king penguins (Aptenodytes patagonicus), which aggregate into large colonies, experience frequent aggressive social interactions, and are regularly exposed to predation by giant petrels (Macronectes sp.) and brown skuas (Catharacta loonbergi) when breeding on land. We approached 200 incubating penguins at four different periods of the breeding season across a range of overall increasing colony densities. We measured the distance at which focal birds detected the approaching threat (alert distance: AD), whether birds decided to flee or not, and the distance of flight initiation (flight initiation distance: FID, viz. the bird attempting to walk away with its egg on its feet). We quantified relative local neighbor density, centrality within the colony (rank), and the number of aggressions the focal bird emitted towards neighbors during the approach. We found that birds engaged in aggressive conflicts with neighbors were less likely to flee, and that increasing relative local neighbor density at low and medium overall colony density resulted in a decrease in bird AD, both supporting the “distracted prey” hypothesis. However, at maximal overall colony density, increasing relative local neighbor density resulted in longer AD, supporting the “many-eyes” hypothesis. We found no support for the “dilution effect” and “selfish herd” hypotheses, and no effects of any hypothesis on FID.

Sex-specific responses to GnRH challenge, but not food supply, in kittiwakes: Evidence for the “sensitivity to information” hypothesis

Seasonal timing of breeding is usually considered to be triggered by endogenous responses linked to predictive cues (e.g., photoperiod) and supplementary cues that vary annually (e.g., food supply), but social cues are also important. Females may be more sensitive to supplementary cues because of their greater role in reproductive timing decisions, while males may only require predictive cues. We tested this hypothesis by food-supplementing female and male colonial seabirds (black-legged kittiwakes, Rissa tridactyla) during the pre-breeding season. We measured colony attendance via GPS devices, quantified pituitary and gonadal responses to gonadotropin releasing hormone (GnRH) challenge, and observed subsequent laying phenology. Food supplementation advanced laying phenology and increased colony attendance. While female pituitary responses to GnRH were consistent across the pre-breeding season, males showed a peak in pituitary sensitivity at approximately the same time that most females were initiating follicle development. The late peak in male pituitary response to GnRH questions a common assumption that males primarily rely on predictive cues (e.g., photoperiod) while females also rely on supplementary cues (e.g., food availability). Instead, male kittiwakes may integrate synchronising cues from their social environment to adjust their reproductive timing to coincide with female timing.

Seabird’s cry: repertoire and vocal expression of contextual valence in the little auk (Alle alle)

Many seabird species breed in colonies counting up to hundreds of thousands of individuals. Life in such crowded colonies might require special coding–decoding systems to reliably convey information through acoustic cues. This can include, for example, developing complex vocal repertoires and adjusting the properties of their vocal signals to communicate behavioural contexts, and thus regulate social interactions with their conspecifics. We studied vocalisations produced by the little auk (Alle alle)—a highly vocal, colonial seabird—over mating and incubation periods on the SW coast of Svalbard. Using passive acoustic recordings registered in a breeding colony, we extracted eight vocalisation types: single call, clucking, classic call, low trill, short call, short-trill, terror, and handling vocalisation. Calls were grouped by production context (based on the typically associated behaviour), to which a valence (positive vs negative) was later attributed, when possible, according to fitness threats, i.e. predator or human presence (negative) and promoters, i.e. interaction with a partner (positive). The effect of the putative valence on eight selected frequency and duration variables was then investigated. The putative contextual valence significantly affected the acoustic properties of the calls. Calls assigned positive valence had higher fundamental frequency and spectral centre of gravity as well as shorter sound duration than these assigned negative valence. These results indicate that the little auk’s vocal communication system may facilitate expression of complex behavioural contexts, and seems to include vocal plasticity within vocalisation types—however, more data are necessary to better understand this effect and possible interplays of other factors.

Sharing your snack: unusual observation of a chick–chick feeding occurrence in colonial king penguin (Aptenodytes patagonicus)

Abstract King penguins (Aptenodytes patagonicus) are colonial seabirds presenting an extraordinary breeding cycle. Parents take over 14 months to raise a single chick to independence, upon which fledglings depart at sea for more than a year. Juveniles often return to the colony within the three austral summers following departure, and before the age of first reproduction (around 5–6 years old), possibly to acquire the essential skills involved in breeding. Little to nothing is known on the acquisition of parental behaviour. Here, we report an anecdotal, yet highly unusual, observation of chick–chick feeding behaviour in this species. The behaviour is highly unusual in that two non-sibling chicks, not yet independent, and hatched at different times (early and late) of the same breeding season were observed, the older chick feeding the younger one. Whereas alloparental feeding is known to occur in this species, this is the first reported observation of a chick–chick feeding event. This unusual behaviour raises the question of whether the early social environment plays a role in the acquisition of essential parenting skills in this species.

Variability, skipped breeding and heavy-tailed dynamics in an Antarctic seabird.

The population dynamics of many colonially breeding seabirds are characterized by large interannual fluctuations that cannot be explained by environmental conditions alone. This variation may be particularly confounded by the use of skipped breeding by seabirds as a life-history strategy, which directly impacts the number of breeding pairs and may affect the accuracy of breeding abundance as a metric of population health. Additionally, large fluctuations in time series may suggest that the underlying population dynamics are heavy tailed, allowing for a higher likelihood of extreme events than expected under Gaussian dynamics. Here, we investigated the effect of demography on time series for abundance of the Adélie penguin Pygoscelis adeliae and explored the occurrence of heavy-tailed dynamics in observed Adélie time series. We focus this study on the Adélie penguin as it is an important bellwether species long used to track the impacts of climate change and fishing on the Southern Ocean ecosystem and shares life-history traits with many colonial seabirds. We quantified the impacts of demographic rates, including skipped breeding, on time series of Adélie abundance simulated using an age-structured model. We also used observed time series of Adélie breeding abundance at all known Antarctic colonies to classify distributions for abundance as Gaussian or non-Gaussian heavy tailed. We then identified the cause of such heavy-tailed dynamics in simulated time series and linked these to spatial patterns in Adélie food resource variability. We found that breeding propensity drives observed breeding fluctuations more than any other vital rate, with high variability in skipped breeding decoupling true abundance from observed breeding abundance. We also found several Antarctic regions characterized by heavy-tailed dynamics in abundance. These regions were often also characterized by high variability in zooplankton availability. In simulated time series, heavy-tailed dynamics were strongly linked to high variability in adult survival. Our results illustrate that stochastic variability in abundance dynamics, particularly the presence of variable rates of skipped breeding, can challenge our interpretation of fluctuations in abundance through time and obscure the relationship between key environmental drivers and population abundance.

Opposing effects of spatiotemporal variation in resources and temporal variation in climate on density dependent population growth in seabirds.

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.

Temporal change in the contribution of immigration to population growth in a wild seabird experiencing rapid population decline

The source–sink paradigm predicts that populations in poorer‐quality habitats (‘sinks') persist due to continued immigration from more‐productive areas (‘sources'). However, this categorisation of populations assumes that habitat quality is fixed through time. Globally, we are in an era of wide‐spread habitat degradation, and consequently there is a pressing need to examine dispersal dynamics in relation to local population change. We used an integrated population model to quantify immigration dynamics in a long‐lived colonial seabird, the black‐legged kittiwakeRissa tridactyla, that is classified as globally ‘Vulnerable'. We then used a transient life table response experiment to evaluate the contribution of temporal variation in vital rates, immigration rates and population structure to realised population growth. Finally, we used a simulation analysis to examine the importance of immigration to population dynamics. We show that the contribution of immigration changed as the population declined. This study demonstrates that immigration is unlikely to maintain vulnerable sink populations indefinitely, emphasising the need for temporal analyses of dispersal to identify shifts that may have dramatic consequences for population viability.

Should I stay or should I go: factors influencing mate retention and divorce in a colonial seabird

Divorce among serially monogamous birds can lead to increased reproductive success if an individual obtains a higher-quality mate or nest site, or it can lead to lower reproductive success due to lack of pair experience or reduced breeding opportunities. Identifying the drivers and consequences of mate retention and divorce are necessary then to understand population dynamics, particularly as animals may alter behaviours in response to climate-induced environmental changes. We used a 33-year data set from a declining Magellanic penguin, Spheniscus magellanicus, colony to identify drivers of mate retention and divorce and test whether the nature of those drivers differed between first-year and long-term pairs. In a pair's first year, breeding outcome was the most important predictor of whether mates would reunite: failure rates were high and 71% of first-year pairs divorced on the whole, but only 47% of pairs that fledged a chick divorced. For long-term pairs, both breeding outcome and between-season changes to nest quality were the most important predictors of whether a pair stayed together: pairs that had a chick hatch or fledge divorced at lower rates than those that failed at the egg stage, and pairs whose nest quality declined were more likely to divorce. We also examined the fitness consequences of divorce by comparing the rates at which females and males obtained new mates and fledged chicks in the following year. Although females were more likely than males to secure a new mate, both sexes experienced similarly increased reproductive success with their new mates compared to their old mates, showing that divorce is adaptive.

Earlier and more frequent occupation of breeding sites during the non-breeding season increases breeding success in a colonial seabird.

Competition for high‐quality breeding sites in colonial species is often intense, such that individuals may invest considerable time in site occupancy even outside the breeding season. The site defense hypothesis predicts that high‐quality sites will be occupied earlier and more frequently, consequently those sites will benefit from earlier and more successful breeding. However, few studies relate non‐breeding season occupancy to subsequent breeding performance limiting our understanding of the potential life‐history benefits of this behavior. Here, we test how site occupancy in the non‐breeding season related to site quality, breeding timing, and breeding success in a population of common guillemots Uria aalge, an abundant and well‐studied colonially breeding seabird. Using time‐lapse photography, we recorded occupancy at breeding sites from October to March over three consecutive non‐breeding seasons. We then monitored the successive breeding timing (lay date) and breeding success at each site. On average, sites were first occupied on the 27th October ± 11.7 days (mean ± SD), subsequently occupied on 46 ± 18% of survey days and for 55 ± 15% of the time when at least one site was occupied. Higher‐quality sites, sites with higher average historic breeding success, were occupied earlier, more frequently and for longer daily durations thereafter. Laying was earlier at sites that were occupied more frequently and sites occupied earlier were more successful, supporting the site defense hypothesis. A path analysis showed that the return date had a greater or equal effect on breeding success as lay date. Pair level occupancy had no effect on breeding timing or success. The clear effect of non‐breeding occupancy of breeding sites on breeding timing and success highlights the benefits of this behavior on demography in this population and the importance of access to breeding sites outside the breeding season in systems where competition for high‐quality sites is intense.

Spatial patterns in age‐ and colony‐specific survival in a long‐lived seabird across 14 contrasting colonies

Demographic rates such as recruitment and survival probability can vary considerably among populations of the same species due to variation in underlying environmental processes. If environmental processes are spatially correlated, nearby populations are expected to have more similar demographic rates than those further apart. Breeding populations and foraging ranges are spatially segregated in colonial seabirds, making them ideal for studying spatial patterns in demographic rates and their effects on local population dynamics. Here we explored variation in age‐dependent survival probabilities across 14 colonies of Herring GullsLarus argentatusbreeding along the Dutch North Sea coast. We used long‐term mark–recapture data of marked fledglings to estimate survival, and estimated spatial autocorrelation of survival probabilities. We assessed whether survival until recruitment age or until 10 years old (close to their expected lifespan) explained variation in population trajectories of each colony. Juvenile and adult survival showed a strong, but different, north‐to‐south gradient in survival probability, with lower juvenile but higher adult survival in northern colonies than southern colonies, whereas the spatial pattern of immature survival was less distinct. Neither recruitment nor the proportion of 10‐year‐old adults alive predicted whether a colony collapsed, declined, remained stable or increased. The distinct spatial pattern in survival suggests variation in regional food availability, which do not seem to drive local population dynamics. The absence of a link between survival and colony trajectories implies that connectivity between populations plays an important role affecting population dynamics.