Individual inter‐annual nest‐site relocation behaviour drives dynamics of a recently established Barnacle Goose Branta leucopsis colony in sub‐arctic Russia

Avian breeding colonies are generally in a continuous state of flux, some parts growing whilst others shrink as individuals move within the colony on the search for better nest‐sites. We examined the role of experience in breeding patch choice by individually marked Barnacle Geese Branta leucopsis in a recently established colony in sub‐arctic Russia. Individuals failing to reproduce successfully tended to shift nest location further the following season than did successful pairs, and they did so towards the most dense nest aggregations within the colony, where reproductive success was higher. We suggest that individual decisions on nest‐site choice shape the spatial dynamics of this colony.     

Influence of density‐dependent competition on foraging and migratory behavior of a subtropical colonial seabird

Density‐dependent competition for food resources influences both foraging ecology and reproduction in a variety of animals. The relationship between colony size, local prey depletion, and reproductive output in colonial central‐place foragers has been extensively studied in seabirds; however, most studies have focused on effects of intraspecific competition during the breeding season, while little is known about whether density‐dependent resource depletion influences individual migratory behavior outside the breeding season. Using breeding colony size as a surrogate for intraspecific resource competition, we tested for effects of colony size on breeding home range, nestling health, and migratory patterns of a nearshore colonial seabird, the brown pelican (Pelecanus occidentalis), originating from seven breeding colonies of varying sizes in the subtropical northern Gulf of Mexico. We found evidence for density‐dependent effects on foraging behavior during the breeding season, as individual foraging areas increased linearly with the number of breeding pairs per colony. Contrary to our predictions, however, nestlings from more numerous colonies with larger foraging ranges did not experience either decreased condition or increased stress. During nonbreeding, individuals from larger colonies were more likely to migrate, and traveled longer distances, than individuals from smaller colonies, indicating that the influence of density‐dependent effects on distribution persists into the nonbreeding period. We also found significant effects of individual physical condition, particularly body size, on migratory behavior, which in combination with colony size suggesting that dominant individuals remain closer to breeding sites during winter. We conclude that density‐dependent competition may be an important driver of both the extent of foraging ranges and the degree of migration exhibited by brown pelicans. However, the effects of density‐dependent competition on breeding success and population regulation remain uncertain in this system.     

Estimating dispersal distributions at multiple scales: within‐colony and among‐colony dispersal rates,distances and directions in European Shags Phalacrocorax aristotelis

Knowledge of the rate, distance and direction of dispersal within and among breeding areas is required to understand and predict demographic and genetic connectivity and resulting population and evolutionary dynamics. However dispersal rates, and the full distributions of dispersal distances and directions, are rarely comprehensively estimated across all spatial scales relevant to wild populations. We used re‐sightings of European Shags Phalacrocorax aristotelis colour‐ringed as chicks on the Isle of May (IoM), UK, to quantify rates, distances and directions of dispersal from natal to subsequent breeding sites both within IoM (within‐colony dispersal) and across 27 other breeding colonies covering 1045 km of coastline (among‐colony dispersal). Additionally, we used non‐breeding season surveys covering 895 km of coastline to estimate breeding season detection probability and hence potential bias in estimated dispersal parameters. Within IoM, 99.6% of individuals dispersed between their natal and observed breeding nest‐site. The distribution of within‐colony dispersal distances was right‐skewed; mean distance was shorter than expected given random settlement within IoM, yet some individuals dispersed long distances within the colony. The distribution of within‐colony dispersal directions was non‐uniform but did not differ from expectation given the spatial arrangement of nest‐sites. However, 10% of all 460 colour‐ringed adults that were located breeding had dispersed to a different colony. The maximum observed dispersal distance (170 km) was much smaller than the maximum distance surveyed (690 km). The distribution of among‐colony dispersal distances was again right‐skewed. Among‐colony dispersal was directional, and differed from random expectation and from the distribution of within‐colony dispersal directions. Non‐breeding season surveys suggested that the probability of detecting a colour‐ringed adult at its breeding location was high in the northeastern UK (98%). Estimated dispersal rates and distributions were therefore robust to incomplete detection. Overall, these data demonstrate skewed and directionally divergent dispersal distributions across small (within‐colony) and large (among‐colony) scales, indicating that dispersal could create genetic and demographic connectivity within the study area.     

The stress hormone corticosterone in a marine top predator reflects short‐term changes in food availability

In many seabird studies, single annual proxies of prey abundance have been used to explain variability in breeding performance, but much more important is probably the timing of prey availability relative to the breeding season when energy demand is at a maximum. Until now, intraseasonal variation in prey availability has been difficult to quantify in seabirds. Using a state‐of‐the‐art ocean drift model of larval cod Gadus morhua, an important constituent of the diet of common guillemots Uria aalge in the southwestern Barents Sea, we were able to show clear, short‐term correlations between food availability and measurements of the stress hormone corticosterone (CORT) in parental guillemots over a 3‐year period (2009–2011). The model allowed the extraction of abundance and size of cod larvae with very high spatial (4 km) and temporal resolutions (1 day) and showed that cod larvae from adjacent northern spawning grounds in Norway were always available near the guillemot breeding colony while those from more distant southerly spawning grounds were less frequent, but larger. The latter arrived in waves whose magnitude and timing, and thus overlap with the guillemot breeding season, varied between years. CORT levels in adult guillemots were lower in birds caught after a week with high frequencies of southern cod larvae. This pattern was restricted to the two years (2009 and 2010) in which southern larvae arrived before the end of the guillemot breeding season. Any such pattern was masked in 2011 by already exceptionally high numbers of cod larvae in the region throughout chick‐rearing period. The findings suggest that CORT levels in breeding birds increase when the arrival of southern sizable larvae does not match the period of peak energy requirements during breeding.     

Among‐colony synchrony in the survival of Common Guillemots Uria aalge reflects shared wintering areas

Spatiotemporal variation in survival may be an important driver of multi‐population dynamics in many wild animal species, yet few scientific studies have addressed this issue, primarily due to a lack of sufficiently comprehensive and detailed datasets. Synchrony in survival rates among different, often distant, subpopulations appears to be common, caused by spatially correlated environmental conditions or by movement of animals from different sites such that their ranges overlap. Many seabird populations are effectively isolated during the breeding season because colonies are widely separated, but over the winter, birds disperse widely and there may be much mixing between different populations. The non‐breeding season is also the period of main mortality for seabirds. Using mark–recapture and ring‐recovery data, we tested for spatial, temporal and age‐related correlations in survival of Common Guillemots Uria aalge among three widely separated Scottish colonies that have varying overlap in their overwintering distributions. Survival was highly correlated over time for colonies/age‐classes sharing wintering areas and, except in 2004, was essentially uncorrelated for those with separate wintering areas. These results strongly suggest that one or more aspects of the winter environment are responsible for spatiotemporal variation in survival of British Guillemots, and provide insight into the factors driving large‐scale population dynamics of the species.     

Small‐scale spatial variation in evolvability for life‐history traits in the storm petrel

The evolutionary potential in the timing of recruitment and reproduction may be crucial for the ability of populations to buffer against environmental changes, allowing them to avoid unfavourable breeding conditions. The evolution of a trait in a local population is determined by its heritability and selection. In the present study, we performed pedigree‐based quantitative genetic analyses for two life‐history traits (recruiting age and laying date) using population data of the storm petrel over an 18‐year period in two adjacent breeding colonies (only 150 m apart) that share the same environmental conditions. In both traits, natal colony effect was the main source of the phenotypic variation among individuals, and cohort variance for recruitment age and additive genetic variance for laying date were natal colony‐specific. We found significant heritability only in laying date and, more specifically, only in birds born in one of the colonies. The difference in genetic variance between the colonies was statistically significant. Interestingly, selection on earlier breeding birds was detected only in the colony in which heritable variation in laying date was found. Therefore, local evolvability for a life‐history trait may vary within a unexpectedly small spatial scale, through the diversifying natural selection and insulating gene flow. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, 106 , 439–446.     

Determinants of between‐year burrow re‐occupation in a colony of the European bee‐eater Merops apiaster

Re‐occupation of existing nesting burrows in the European bee‐eater Merops apiaster has only rarely – and if so mostly anecdotically – been documented in the literature record, although such behavior would substantially save time and energy. In this study, we quantify burrow re‐occupation in a German colony over a period of eleven years and identify ecological variables determining reuse probability. Of 179 recorded broods, 54% took place in a reused burrow and the overall probability that one of 75 individually recognized burrows would be reused in a given subsequent year was estimated as 26.4%. This indicates that between‐year burrow reuse is a common behavior in the study colony which contrasts with findings from studies in other colonies. Furthermore, burrow re‐occupation probability declined highly significantly with increasing age of the breeding wall. Statistical separation of within‐ and between‐burrow effects of the age of the breeding wall revealed that a decline in re‐occupation probability with individual burrow age was responsible for this and not a selective disappearance of burrows with high re‐occupation probability over time. Limited duty cycles of individual burrows may be caused by accumulating detritus or decreasing stability with increasing burrow age. Alternatively, burrow fidelity may presuppose pair fidelity which may also explain the observed restricted burrow reuse duty cycles. A consequent next step would be to extend our within‐colony approach to other colonies and compare the ecological circumstances under which bee‐eaters reuse breeding burrows.     

Visual tracking from a rigid‐hulled inflatable boat to determine foraging movements of breeding terns

ABSTRACT Defining the at‐sea foraging movements of seabirds is fundamental to understanding their ecology and can also be important in assessing the potential impact of marine developments such as offshore wind farms (OWFs). Surveys of predefined areas using aerial or boat‐based transect surveys may not allow adequate assessment of the relative importance of different areas to birds. Individual‐based satellite or radio‐telemetry can be effective in identifying foraging ranges and preferred areas, but may not be suitable for some species. We developed a method to determine the foraging movements of breeding terns (Sterna spp.) by visually tracking individuals using a rigid‐hulled inflatable boat (RHIB). Sandwich Terns (S. sandvicensis), Common Terns (S. hirundo), and Arctic Terns (S. paradisaea) were tracked from colonies in Norfolk and Anglesey, United Kingdom, from 2006 to 2008. The proportion of complete (from and to colony) trips varied from 29–60% among species, years, and colonies. Individual Sandwich Terns were tracked for periods up to 126 min over distances up to 72 km and as far as 54 km from the breeding colony, further than Arctic (up to 57 km and 29 km from the colony) and Common (to 29 km and <9 km from the colony) terns. Mean values were much smaller. Multivariate modeling of Sandwich Tern foraging trips indicated that flight speeds >50 km/hr coupled with greater distances from shore (>25 km) significantly reduced the likelihood of tracking a bird for an entire foraging trip. Use of different boats that differ in speed and performance may alleviate such issues. Visual tracking allowed us to collect data on foraging behavior, flight height, and prey capture rates, and also permitted comparisons between species. Our results indicate that visual tracking may be an effective means of determining the foraging movements and at‐sea behavior of a variety of short‐ranging, day‐active seabirds.     

Inter‐colony foraging area segregation quantified in small colonies of Adélie Penguins

Theoretical models on the movement of colonial animals predict that neighbouring colonies may segregate their foraging areas, and many seabird studies have reported the presence of such segregations. However, these studies have often lacked the appropriate null model to test the effect of neighbouring colonies on foraging areas, especially in small colonies or in short‐ranging species. Here, we examined the foraging areas of Adélie Penguins Pygoscelis adeliae from two neighbouring (2 km apart) colonies by using bird‐borne GPS loggers. The field study was conducted at Hukuro Cove colony (104 pairs) and Mizukuguri Cove colony (338 pairs) in Lützow‐Holm Bay, East Antarctica. We obtained GPS tracks for 504 foraging trips from 48 chick‐rearing Adélie Penguins and quantified the degree of overlap in the foraging areas between two colonies. We also produced simulated movement tracks by using correlated random‐walks assuming no inter‐colony competition and quantified the degree of overlap in the simulated foraging areas. Finally, we compared the results from real GPS tracks with those from simulated tracks to examine the effect of neighbouring colonies on Adélie Penguin movement. The results indicate that the degree of overlap was significantly smaller in real tracks than in simulated tracks. In real tracks, the foraging area of the smaller Hukuro Cove colony extended to the other side of the larger Mizukuguri Cove colony, unlike in simulated tracks. Consequently, we suggest that Adélie Penguins from two neighbouring colonies segregated their foraging areas and that the larger colony appeared to affect the foraging area of the smaller colony.     

Seasonal dynamics of agonistic behavior and hormones in an ex situ all‐male colony of large flying foxes

Large flying foxes (Pteropus vampyrus) are a socially complex species. In situ colonies typically comprise thousands of individuals in small harems of one male to many females. In ex situ environments, all‐male colonies are becoming more common due to a surplus of males in the population. There is limited information describing the hormonal and behavioral patterns of all‐male colonies during the breeding season. We assessed seasonal changes in hormones and behavior in an all‐male colony of 12 large flying foxes at Disney's Animal Kingdom^®. We validated hormone assays using morning urine and fecal samples to assess seasonal changes in excreted immunoreactive testosterone and glucocorticoid metabolites. We collected behavior data using an all‐occurrence method, recording agonistic behaviors related to territorial defense (hooking, biting, wing flexing, vocalizing, and wrestling), and sexual behavior (mounting and frontal grabbing). Results indicated that (i) we could reliably measure testosterone and glucocorticoid metabolites concentrations from fecal and urine samples collected from individual bats; (ii) there were distinct relationships between changes in levels of agonism and hormone concentrations throughout the year; and (iii) three agonistic behaviors (chasing, wrestling, and open‐mouth threat) peaked prior to the increase in testosterone and glucocorticoid hormones measured during the breeding season. These three behaviors could potentially be used as early indicators to signal the onset of the breeding season and allow time to implement ex situ management changes to reduce the incidence of agonism between individuals.     

High foraging site fidelity and spatial segregation among individual great black‐backed gulls

Individual foraging site fidelity, whereby individuals repeatedly visit the same foraging areas, is widespread in nature, and likely benefits individuals through higher foraging efficiency and potentially, higher breeding success. It may arise as a consequence of habitat or resource specialisation, or alternatively, where resources are abundant or predictable, the partitioning of space might guarantee individuals exclusive foraging opportunities. We tracked seven adult great black‐backed gulls Larus marinus at a North Sea colony from early incubation to the end of the breeding season in 2016, providing a total of 1170 foraging trips over a mean ± SD tracking period of 67 ± 16 days. There was clear spatial segregation between individuals, with almost no overlap of their core areas (50% utilisation distribution) during incubation and chick‐rearing. Core areas were relatively small and there was high repeatability (R ± SE) in foraging parameters, including initial departure direction (0.73 ± 0.11), foraging range (0.41 ± 0.14) and cumulative distance travelled (0.19 ± 0.1) throughout the breeding season. Despite the low spatial overlap, there was little evidence of differential habitat use by individuals. The near‐exclusive individual foraging areas of this species, usually considered to be a generalist, indicate that where there is high resource availability throughout the breeding season and a small local population, individuals appear to adopt a territorial strategy which likely reduces intraspecific competition.     

Dispersal and recruitment during population growth in a colonial bird, the great cormorant Phalacrocorax carbo sinensis

While the factors influencing reproduction and survival in colonial populations are relatively well studied, factors involved in dispersal and settlement decisions are not well understood. The present study investigated exchanges of great cormorants Phalacrocorax carbo sinensis among six breeding colonies over a 13‐year period when the breeding population in Denmark increased from 2800 to 36 400 nests. We used a multistate capture‐recapture model that combined multisite resightings and recoveries to examine simultaneously recruitment, natal dispersal, breeding dispersal and annual survival of first‐year, immature and breeding great cormorants. Mean survival of first‐year birds (0.50±0.09, range=0.42–0.66 among colonies) was lower than survival of breeders (0.90±0.06, range=0.81–0.97). Mean survival of immature birds over the study period was 0.87±0.08. Dispersal from a colony increased with decreasing mean brood size in the colony in both first‐time and experienced breeders. The choice of the settlement colony in first‐time breeders was affected by conditions in the natal colony and in the colonies prospected during the pre‐breeding years. In particular, first‐time breeders recruited to colonies where they could expect better breeding success. Experienced breeders relied mainly on cues present early in the season and on their own breeding experience to choose a new breeding colony. Newly established colonies resulted mainly from the immigration of first‐time breeders originating from denser colonies. Dispersal was distance‐dependent and first‐time breeders dispersed longer distances than breeders. We suggest that the prospecting behaviour allows first‐time breeders to recruit in nearby as well as more distant potential breeding colonies. Dispersing breeders preferred to settle in neighbouring colonies likely to benefit from their experience with foraging areas. We discuss the importance of these movements for growth and expansion of the breeding population.     

Recruitment to a seabird population depends on environmental factors and on population size

1. A novel capture-mark-recapture (CMR) method was used to build a multistate model of recruitment by young birds to a breeding population of common guillemots Uria aalge on the Isle of May, Scotland. Recruitment of a total of 2757 individually marked guillemots over 17 years was modelled as a process where individuals had to move from an unobservable state at sea, through a nonbreeding state present in the colony, to the breeding state. The probabilities of individuals returning to the colony in a given year, at age 2 and 3-4 years, were positively correlated with an environmental covariate, the winter North Atlantic Oscillation index (WNAO) in the previous years. 2. For 2 year olds, there was a negative relationship with breeding population size, suggesting that density dependence operated in this colony through limitation of food or some other resource. 3. Survival over the first 2 years of life varied with cohort, but was unrelated to the WNAO. Mean survival over this 2-year period was high at 0.576 (95% CI: 0.444; 0.708). 4. This high survival, combined with a low 'local' survival after age 5 years of 0.695 (0-654; 0.733) and observations of Isle of May chicks at other colonies, suggests that most surviving chicks return to the natal colony before deciding whether to recruit there or move elsewhere.     

Multicolony tracking reveals the winter distribution of a pelagic seabird on an ocean basin scale

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.     

Fine‐scale impacts on avian biodiversity due to a despotic species,the bell miner (Manorina melanophrys)

Bell miners (Manorina melanophrys; Meliphagidae) are a highly social and very aggressive honeyeater. They are despotic and cooperate in the defence of their territories against other bird species, leading to the almost complete exclusion of other avifauna from miner‐occupied regions. This study aimed to resolve some of the fine‐scale effects of bell miner aggression on avian diversity both within and adjacent to colonies to determine the true impact of a colony on local avifaunal abundance. Three areas, distributed throughout the range of the bell miner, were surveyed across both non‐breeding and breeding seasons to assess the temporal and spatial impacts of bell miner aggression on other bird species. Bell miner colonies were found to occupy very clearly defined areas and had the expected negative impact on avian diversity within their colony. The effects of bell miner colony presence on abundance and richness of avian species were found to cease at the colony boundary, with both recovering to normal levels immediately outside the bell miner colony. Whether bell miners were breeding or not, and irrespective of the amount of vegetation coverage, bell miner colonies were found to have relatively marginal impacts on avian richness and abundance. No impact of colony presence/absence was found on the richness or abundance of the avian species that dwell in the undergrowth, with some evidence that these species were actually more common at the colony edge. Our results demonstrate that the influence of bell miner colony presence upon avian biodiversity is restricted to the confines of the colony and does not radiate outwards into the surrounding habitat. Colony presence influences, therefore, have implications when considering the impact of bell miner behaviour on the diversity of insectivorous birds and processes, most notably the propagation of Bell Miner Associated Dieback.     

Distribution of the bridled form of the Common guillemot Uria aalge in the North Atlantic

This study documents the fourth survey (198 1–1982) of bridled guillemots and provides the most complete information yet available on the distribution of the bridled form of the Common guillemot. The distribution has remained largely unchanged since the last survey (1959-60), although marked changes have occurred at a few colonies. Sea surface temperature "explains" only 35% of the variance in the proportion of bridled guillemots. Minimum air temperatures during the breeding season are much more closely correlated with bridling and "explain" 60% of the variance. No relationship exists between sea temperatures and the proportion of bridled guillemots in wintering areas. The distribution of bridled guillemots may represent the slow, southward spread of the bridled form. Alternatively, selection may favour the bridled form in cooler areas. There is insufficient evidence to distinguish between these two hypotheses, but if selection is important, the correlations with environmental factors suggests that it operates during the breeding season rather than at other times.     

Species‐specific foraging strategies and segregation mechanisms of sympatric Antarctic fulmarine petrels throughout the annual cycle

Determining the year‐round distribution and behaviour of birds is necessary for a better understanding of their ecology and foraging strategies. Petrels form an important component of the high‐latitude seabird assemblages in terms of species and individuals. The distribution and foraging ecology of three sympatric fulmarine petrels (Southern Fulmar Fulmarus glacialoides, Cape Petrel Daption capense and Snow Petrel Pagodroma nivea) were studied at Adélie Land, East Antarctica, by combining information from miniaturized saltwater immersion geolocators and stable isotopes from feathers. During the breeding season at a large spatial scale (c. 200 km), the three species overlapped in their foraging areas located in the vicinity of the colonies but were segregated by their diet and trophic level, as indicated by the different chick δ¹⁵N values that increased in the order Cape Petrel < Southern Fulmar < Snow Petrel. During the non‐breeding season, the three fulmarines showed species‐specific migration strategies along a wide latitudinal gradient. Snow Petrels largely remained in ice‐associated Antarctic waters, Southern Fulmars targeted primarily the sub‐Antarctic zone and Cape Petrels migrated further north. Overall, birds spent less time in flight during the non‐breeding period than during the breeding season, with the highest percentage of time spent sitting on the water occurring during the breeding season and at the beginning of the non‐breeding period before migration. This activity pattern, together with the δ¹³C values of most feathers, strongly suggests that moult of the three fulmarine petrels occurred at that time in the very productive high Antarctic waters, where birds fed on a combination of crustaceans and fish. The study highlights different segregating mechanisms that allow the coexistence of closely related species, specifically, prey partitioning during the breeding season and spatial segregation at sea during the non‐breeding season.     

Intercolony variation in foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period

Using GPS loggers, we examined the influence of colony, sex, and bird identity on foraging flight characteristics of black‐headed gulls Chroicocephalus ridibundus during the incubation period. We studied tracks of 36 individuals breeding in one urban and two rural colonies in Poland. Birds from both rural colonies performed the furthest flights (mean max distance 8–12 km, up to 27 km) foraging mainly in agricultural areas. Gulls from the urban colony performed shorter flights (mean 5 km, up to 17 km) visiting mainly urbanized areas and water bodies. We found that females performed longer flights and their flight parameters were less repeatable compared to males. Males from both rural colonies visited water bodies more frequently than females. In all colonies, males (but not females) used habitats unproportionally to their availability in the vicinity. Relatively low interindividual and relatively high intraindividual overlap in home ranges indicated considerable foraging site fidelity. Individuals specialized in the use of a particular type of habitat performed shorter foraging flights compared to individuals using diverse habitats during their foraging flights. Our results indicate diverse foraging strategies of black‐headed gulls, including generalists that explore various habitats and specialists characterized by high foraging site and habitat fidelity.     

Strong temporal consistency in the individual foraging behaviour of Imperial Shags Phalacrocorax atriceps

Individual consistency in foraging behaviour can generate behavioural variability within populations and may, ultimately, lead to species diversification. However, individual‐based long‐term behavioural studies are particularly scarce in seabird species. Between 2008 and 2011, breeding Imperial Shags Phalacrocorax atriceps at the Punta León colony, Argentina, were tracked with GPS devices to evaluate behavioural consistency during their foraging trips. Within a breeding season, individuals were highly consistent in the maximum distances they reached from the shore and the colony, as well as in the time invested in flight and diving across consecutive days during early chick rearing. In addition, each individual had its specific foraging area distinct from the foraging area of other individuals. Comparing between early and late chick rearing in the same season, individuals were consistent, to a lesser degree, in the maximum distance they reached from the colony and the shore, increasing in consistency later on in the season. Within the season, females were more consistent than males in the maximum distance they moved from the colony and the shore, the sexes segregated in their foraging areas and individual females were segregated from one another. Twenty‐eight individuals tracked in different breeding seasons were marginally consistent in their trip durations and maximum distance reached from shore across seasons. Among seasons, foraging locations differed between sexes and among individual females. Individuals from this colony exhibited consistency over time in several aspects of foraging behaviour, which may be due to a combination of individual characteristics such as learning abilities, breeding experience or health, as well as targeted prey type and stability of the environment at this location.     

Central–periphery gradient of individual quality within a colony of Black‐headed Gulls

Central nesting sites within avian colonies are often more profitable in terms of fitness, as they can offer better protection against predators compared with colony edges. Thus, central sites are expected to attract high‐quality individuals, which should produce a clear central–periphery gradient in the phenotypic quality of nesting individuals and reproductive output within colonies. The aim of this study was to assess spatial patterns of individual quality within a colony of a common larid, the Black‐headed Gull Chroicocephalus ridibundus. We found that laying was advanced in the central zone of the colony when compared with the periphery, and central pairs tended to lay larger eggs and had higher hatching success than peripheral pairs. Centrally nesting individuals had higher blood haemoglobin concentrations, size‐corrected body mass and heterozygosity (males only), indicating their better condition and higher genetic quality. In contrast, central females had lower plasma albumin concentration, which could possibly be attributed to their greater investment in egg production. Finally, pairs breeding in the central zone of the colony had an elevated heterophil/lymphocyte ratio, suggesting that social stress can be recognized as an important cost of nesting in dense aggregations. The results provide empirical support for central–periphery gradients in the individual quality of colonially nesting Black‐headed Gulls, indicating that spatial gradients in pair quality should be carefully taken into account when conducting research on colonial birds.