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.     

Diet segregation between two colonies of little penguins Eudyptula minor in southeast Australia

We studied foraging segregation between two different sized colonies of little penguins Eudyptula minor with overlapping foraging areas in pre‐laying and incubation. We used stomach contents and stable isotope measurements of nitrogen (δ¹⁵N) and carbon (δ¹³C) in blood to examine differences in trophic position, prey‐size and nutritional values between the two colonies. Diet of little penguins at St Kilda (small colony) relied heavily on anchovy while at Phillip Island (large colony), the diet was more diverse and anchovies were larger than those consumed by St Kilda penguins. Higher δ¹⁵N values at St Kilda, differences in δ¹³C values and the prey composition provided further evidence of diet segregation between colonies. Penguins from each colony took anchovies from different cohorts and probably different stocks, although these sites are only 70 km apart. Differences in diet were not reflected in protein levels in the blood of penguins, suggesting that variation in prey between colonies was not related to differences in nutritional value of the diet. Anchovy is currently the only available prey to penguins throughout the year and its absence could have a negative impact on penguin food supply, particularly at St Kilda where the diet is dominated by this species. While it is difficult to establish whether diet segregation is caused by inter‐ or intra‐colony competition or spatial differences in foraging areas, we have shown that colonies with broadly overlapping foraging ranges could have significant differences in trophic position, diet composition and prey size while maintaining a diet of similar nutritional value.     

Differential adult survival at close seabird colonies: The importance of spatial foraging segregation and bycatch risk during the breeding season

Marine megafauna, including seabirds, are critically affected by fisheries bycatch. However, bycatch risk may differ on temporal and spatial scales due to the uneven distribution and effort of fleets operating different fishing gear, and to focal species distribution and foraging behavior. Scopoli's shearwater Calonectris diomedea is a long‐lived seabird that experiences high bycatch rates in longline fisheries and strong population‐level impacts due to this type of anthropogenic mortality. Analyzing a long‐term dataset on individual monitoring, we compared adult survival (by means of multi‐event capture–recapture models) among three close predator‐free Mediterranean colonies of the species. Unexpectedly for a long‐lived organism, adult survival varied among colonies. We explored potential causes of this differential survival by (1) measuring egg volume as a proxy of food availability and parental condition; (2) building a specific longline bycatch risk map for the species; and (3) assessing the distribution patterns of breeding birds from the three study colonies via GPS tracking. Egg volume was very similar between colonies over time, suggesting that environmental variability related to habitat foraging suitability was not the main cause of differential survival. On the other hand, differences in foraging movements among individuals from the three colonies expose them to differential mortality risk, which likely influenced the observed differences in adult survival. The overlap of information obtained by the generation of specific bycatch risk maps, the quantification of population demographic parameters, and the foraging spatial analysis should inform managers about differential sensitivity to the anthropogenic impact at mesoscale level and guide decisions depending on the spatial configuration of local populations. The approach would apply and should be considered in any species where foraging distribution is colony‐specific and mortality risk varies spatially.     

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.     

Colony size and foraging range in seabirds

The reasons for variation in group size among animal species remain poorly understood. Using ‘Ashmole's halo’ hypothesis of food depletion around colonies, we predict that foraging range imposes a ceiling on the maximum colony size of seabird species. We tested this with a phylogenetic comparative study of 43 species of seabirds (28 262 colonies), and investigated the interspecific correlation between colony size and foraging ranges. Foraging range showed weak relationships with the low percentiles of colony size of species, but the strength of the association increased for larger percentiles, peaking at the maximum colony sizes. To model constraints on the functional relationship between the focal traits, we applied a quantile regression based on maximum colony size. This showed that foraging range imposes a constraint to species’ maximum colony sizes with a slope around 2. This second‐order relationship is expected from the equation of the area of a circle. Thus, our large dataset and innovative statistical approach shows that foraging range imposes a ceiling on seabird colony sizes, providing strong support to the hypothesis that food availability is an important regulator of seabird populations.     

Taking a trip to the shelf: Behavioral decisions are mediated by the proximity to foraging habitats in the black‐legged kittiwake

For marine top predators like seabirds, the oceans represent a multitude of habitats regarding oceanographic conditions and food availability. Worldwide, these marine habitats are being altered by changes in climate and increased anthropogenic impact. This is causing a growing concern on how seabird populations might adapt to these changes. Understanding how seabird populations respond to fluctuating environmental conditions and to what extent behavioral flexibility can buffer variations in food availability can help predict how seabirds may cope with changes in the marine environment. Such knowledge is important to implement proper long‐term conservation measures intended to protect marine predators. We explored behavioral flexibility in choice of foraging habitat of chick‐rearing black‐legged kittiwakes Rissa tridactyla during multiple years. By comparing foraging behavior of individuals from two colonies with large differences in oceanographic conditions and distances to predictable feeding areas at the Norwegian shelf break, we investigated how foraging decisions are related to intrinsic and extrinsic factors. We found that proximity to the shelf break determined which factors drove the decision to forage there. At the colony near the shelf break, time of departure from the colony and wind speed were most important in driving the choice of habitat. At the colony farther from the shelf break, the decision to forage there was driven by adult body condition. Birds furthermore adjusted foraging behavior metrics according to time of the day, weather conditions, body condition, and the age of the chicks. The study shows that kittiwakes have high degree of flexibility in their behavioral response to a variable marine environment, which might help them buffer changes in prey distribution around the colonies. The flexibility is, however, dependent on the availability of foraging habitats near the 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.     

Assessing the effectiveness of foraging radius models for seabird distributions using biotelemetry and survey data

Relatively simple foraging radius models have the potential to generate predictive distributions for a large number of species rapidly, thus providing a cost-effective alternative to large-scale surveys or complex modelling approaches. Their effectiveness, however, remains largely untested. Here we compare foraging radius distribution models for all breeding seabirds in Ireland, to distributions of empirical data collected from tracking studies and aerial surveys. At the local/colony level, we compared foraging radius distributions to GPS tracking data from seabirds with short (Atlantic puffin Fratercula arctica, and razorbill Alca torda) and long (Manx shearwater Puffinus puffinus, and European storm-petrel Hydrobates pelagicus) foraging ranges. At the regional/national level, we compared foraging radius distributions to extensive aerial surveys conducted over a two-year period. Foraging radius distributions were significantly positively correlated with tracking data for all species except Manx shearwater. Correlations between foraging radius distributions and aerial survey data were also significant, but generally weaker than those for tracking data. Correlations between foraging radius distributions and aerial survey data were benchmarked against generalised additive models (GAMs) of the aerial survey data that included a range of environmental covariates. While GAM distributions had slightly higher correlations with aerial survey data, the results highlight that the foraging radius approach can be a useful and pragmatic approach for assessing breeding distributions for many seabird species. The approach is likely to have acceptable utility in complex, temporally variable ecosystems and when logistic and financial resources are limited.     

Climate oscillation and the invasion of alien species influence the oceanic distribution of seabirds

Spatial and temporal distribution of seabird transiting and foraging at sea is an important consideration for marine conservation planning. Using at‐sea observations of seabirds (n = 317), collected during the breeding season from 2012 to 2016, we built boosted regression tree (BRT) models to identify relationships between numerically dominant seabird species (red‐footed booby, brown noddy, white tern, and wedge‐tailed shearwater), geomorphology, oceanographic variability, and climate oscillation in the Chagos Archipelago. We documented positive relationships between red‐footed booby and wedge‐tailed shearwater abundance with the strength in the Indian Ocean Dipole, as represented by the Dipole Mode Index (6.7% and 23.7% contribution, respectively). The abundance of red‐footed boobies, brown noddies, and white terns declined abruptly with greater distance to island (17.6%, 34.1%, and 41.1% contribution, respectively). We further quantified the effects of proximity to rat‐free and rat‐invaded islands on seabird distribution at sea and identified breaking point distribution thresholds. We detected areas of increased abundance at sea and habitat use‐age under a scenario where rats are eradicated from invaded nearby islands and recolonized by seabirds. Following rat eradication, abundance at sea of red‐footed booby, brown noddy, and white terns increased by 14%, 17%, and 3%, respectively, with no important increase detected for shearwaters. Our results have implication for seabird conservation and island restoration. Climate oscillations may cause shifts in seabird distribution, possibly through changes in regional productivity and prey distribution. Invasive species eradications and subsequent island recolonization can lead to greater access for seabirds to areas at sea, due to increased foraging or transiting through, potentially leading to distribution gains and increased competition. Our approach predicting distribution after successful eradications enables anticipatory threat mitigation in these areas, minimizing competition between colonies and thereby maximizing the risk of success and the conservation impact of eradication programs.     

Genetic divergence between colonies of Flesh-footed Shearwater <Emphasis Type="Italic">Ardenna carneipes</Emphasis> exhibiting different foraging strategies

Increasing evidence suggests foraging segregation as a key mechanism promoting genetic divergence within seabird species. However, testing for a relationship between population genetic structure and foraging movements among seabird colonies can be challenging. Telemetry studies suggest that Flesh-footed Shearwater Ardenna carneipes that breed at Lord Howe Island or New Zealand, versus southwestern Australia or Saint-Paul Island in the Indian Ocean, migrate to different regions (North Pacific Ocean and northern Indian Ocean, respectively) during the non-breeding season, which may inhibit gene flow among colonies. In this study, we sequenced a 858-base pair mitochondrial region and seven nuclear DNA fragments (352–654 bp) for 148 individuals to test genetic differentiation among colonies of Flesh-footed Shearwaters. Strong genetic divergence was detected between Pacific colonies relative to those further West. Molecular analysis of fisheries’ bycatch individuals sampled in the Sea of Japan indicated that individuals from both western and eastern colonies were migrating through this area, and hence the apparent segregation of the non-breeding distribution based on telemetry is invalid and cannot contribute to the population genetic structure among colonies. The genetic divergence among colonies is better explained by philopatry and evidence of differences in foraging strategies during the breeding season, as supported by the observed genetic divergence between Lord Howe Island and New Zealand colonies. We suggest molecular analysis of fisheries’ bycatch individuals as a rigorous method to identify foraging segregation, and we recommend the eastern and western A. carneipes colonies be regarded as different Management Units.     

An energetic correlate between colony size and foraging effort in seabirds, an example of the Adélie penguin Pygoscelis adeliae

Central-place foraging seabirds alter the availability of their prey around colonies, forming a "halo" of reduced prey access that ultimately constrains population size. This has been indicated indirectly by an inverse correlation between colony size and reproductive success, numbers of conspecifics at other colonies within foraging range, foraging effort (i.e. trip duration), diet quality and colony growth rate. Although ultimately mediated by density dependence relative to food through intraspecific exploitative or interference competition, the proximate mechanism involved has yet to be elucidated. Herein, we show that Adélie penguin Pygoscelis adeliae colony size positively correlates to foraging trip duration and metabolic rate, that the metabolic rate while foraging may be approaching an energetic ceiling for birds at the largest colonies, and that total energy expended increases with trip duration although uncompensated by increased mass gain. We propose that a competition-induced reduction in prey availability results in higher energy expenditure for birds foraging in the halo around large colonies, and that to escape the halo a bird must increase its foraging distance. Ultimately, the total energetic cost of a trip determines the maximum successful trip distance, as on longer trips food acquired is used more for self maintenance than for chick provisioning. When the net cost of foraging trips becomes too high, with chicks receiving insufficient food, chick survival suffers and subsequent colony growth is limited. Though the existence of energetic studies of the same species at multiple colonies is rare, because foraging metabolic rate increases with colony size in at least two other seabird species, we suggest that an energetic constraint to colony size may generally apply to other seabirds.     

Varying foraging patterns in response to competition? A multicolony approach in a generalist seabird

Reducing resource competition is a crucial requirement for colonial seabirds to ensure adequate self‐ and chick‐provisioning during breeding season. Spatial segregation is a common avoidance strategy among and within species from neighboring breeding colonies. We determined whether the foraging behaviors of incubating lesser black‐backed gulls (Larus fuscus) differed between six colonies varying in size and distance to mainland, and whether any differences could be related to the foraging habitats visited. Seventy‐nine incubating individuals from six study colonies along the German North Sea coast were equipped with GPS data loggers in multiple years. Dietary information was gained by sampling food pellets, and blood samples were taken for stable isotope analyses. Foraging patterns clearly differed among and within colonies. Foraging range increased with increasing colony size and decreased with increasing colony distance from the mainland, although the latter might be due to the inclusion of the only offshore colony. Gulls from larger colonies with consequently greater density‐dependent competition were more likely to forage at land instead of at sea. The diets of the gulls from the colonies furthest from each other differed, while the diets from the other colonies overlapped with each other. The spatial segregation and dietary similarities suggest that lesser black‐backed gulls foraged at different sites and utilized two main habitat types, although these were similar across foraging areas for all colonies except the single offshore island. The avoidance of intraspecific competition results in colony‐specific foraging patterns, potentially causing more intensive utilization of terrestrial foraging sites, which may offer more predictable and easily available foraging compared with the marine environment.     

Physiological effects of climate on distributions of endothermic species

Aim Determining the mechanisms underlying climatic limitation of species distributions is essential for understanding responses to current climatic change. Disentangling direct (e.g. physiological) and indirect (e.g. trophic) effects of climate on distributions through occurrence‐based modelling is problematic because most species use the same area for both shelter and food acquisition. By focusing on marine birds that breed on land but feed at sea, we exploit a rare opportunity to dissociate direct from indirect climatic effects on endothermic species. Location Coastal Europe. Methods We developed climate‐response surfaces (CRS) for 13 seabird species in coastal Europe, linking terrestrial climatic variables considered important for heat transfer with presence/absence data across each species’ entire European breeding range. Agreement between modelled and actual distribution was assessed for jackknifed samples using area under the curve (AUC) of receiver operating characteristic plots. Higher AUC values indicated closer correspondence between observed breeding distribution and terrestrial climate. We assessed the influence of several ecological factors on model performance across species. Results Species maximum foraging range and breeding latitude explained the greatest proportion of variation in AUC across species. AUC was positively related to both latitude and foraging range. Main conclusions The positive relationship between foraging range and AUC suggests that species foraging further are more likely to be constrained by environmental heat stress conditions at the breeding site. One plausible explanation is that long foraging trips result in one parent spending long periods in continuous nest attendance, exposed to such conditions. These may include negative impacts through predation and parasitism in addition to physiological responses to the thermal environment, which probably explains why our models performed better for species breeding at higher latitudes, where such species interactions are considered less important. These data highlight the importance of considering physiological impacts of climate for endothermic species, and suggest that widespread oceanographic changes that reduce prey quality and quantity for seabirds at sea may be exacerbated by additional impacts of climate at the breeding site.     

At‐sea distribution and habitat use in king penguins at sub‐Antarctic Marion Island

King penguins make up the bulk of avian biomass on a number of sub‐Antarctic islands where they have a large functional effect on terrestrial and marine ecosystems. The same applies at Marion Island where a substantial proportion of the world population breeds. In spite of their obvious ecological importance, the at‐sea distribution and behavior of this population has until recently remained entirely unknown. In addressing this information deficiency, we deployed satellite‐linked tracking instruments on 15 adult king penguins over 2 years, April 2008 and 2013, to study their post‐guard foraging distribution and habitat preferences. Uniquely among adult king penguins, individuals by and large headed out against the prevailing Antarctic Circumpolar Current, foraging to the west and southwest of the island. On average, individuals ventured a maximum distance of 1,600 km from the colony, with three individuals foraging close to, or beyond, 3,500 km west of the colony. Birds were mostly foraging south of the Antarctic Polar Front and north of the southern boundary of the Antarctic Circumpolar Current. Habitat preferences were assessed using boosted regression tree models which indicated sea surface temperate, depth, and chorophyll a concentration to be the most important predictors of habitat selection. Interestingly, king penguins rapidly transited the eddy‐rich area to the west of Marion Island, associated with the Southwest Indian Ocean Ridge, which has been shown to be important for foraging in other marine top predators. In accordance with this, the king penguins generally avoided areas with high eddy kinetic energy. The results from this first study into the behavioral ecology and at‐sea distribution of king penguins at Marion Island contribute to our broader understanding of this species.     

Balancing food and density‐dependence in the spatial distribution of an interference‐prone forager

Foraging distributions are thought to be density‐dependent, because animals not only select for a high availability and quality of resources, but also avoid conspecific interference. Since these processes are confounded, their relative importance in shaping foraging distributions remains poorly understood. Here we aimed to rank the contribution of density‐dependent and density‐independent effects on the spatio‐temporal foraging patterns of eurasian oystercatchers. In our intertidal study area, tides caused continuous variation in oystercatcher density, providing an opportunity to disentangle conspecific interference and density‐independent interactions with the food landscape. Spatial distributions were quantified using high‐resolution individual tracking of foraging activity and location. In a model environment that included a realistic reconstruction of both the tides and the benthic food, we tested a family of behaviour‐based optimality models against these tracking data. Density‐independent interactions affected spatial distributions much more strongly than conspecific interference, even in an interference‐prone species like oystercatchers. Spatial distributions were governed by avoidance of bill injury costs, selection for high interference‐free intake rates and a decreasing availability of benthic bivalve prey after their exposure. These density‐independent interactions outweighed interference competition in terms of effect size. We suggest that the bottleneck in our mechanistic understanding of foraging distributions may be primarily the role of density‐independent prey attributes unrelated to intake rates, like damage costs in the case of oystercatchers foraging on perilous prey. At a landscape scale, above the finest inter‐individual distances, effects of conspecific interaction on spatial distributions may have been overemphasised.     

Colony size and not nest density drives reproductive output in the Common Tern Sterna hirundo

Density is known to be an important factor in population size regulation. Several mechanisms of density limitation have been identified in colonial birds. We studied competition in Common Terns Sterna hirundo to assess whether the factor limiting reproductive output was competition for nest‐sites, which is dependent on local nest density, or density‐dependent competition for food resources, which is dependent on overall colony size using the same foraging area. We found strong associations of both colony size and nest density with reproductive output in five colonies of Common Terns in three different habitats (one marine, two freshwater). Based on detailed long‐term datasets of six separate sub‐colonies of the Banter See colony that differed in nest density, we found that reproductive success was not related to nest density but to overall colony size, possibly a result of resource depletion and food competition. We also found carry‐over effects of colony size during rearing on post‐fledging return rate. These results have important implications for the conservation management plans aimed at recovering declining populations of Common Terns.     

Spatial segregation between immatures and adults in a pelagic seabird suggests age‐related competition

Individual competitiveness conditions access to resources when they are limited. Immature individuals that are less skilled than adults have to adapt their foraging strategies to survive. Among strategies to reduce competition, spatial segregation has been widely demonstrated. However, the use of spatial segregation by immatures to limit intra‐specific competition with adults has rarely been tested. In this study, we investigated and compared habitat preferences and distributions of free‐ranging immature and breeding adult northern gannets Morus bassanus in order to determine whether they compete for similar habitats during the year, and if this results in a spatial segregation between birds of different age groups. Based on > 66 000 km of aerial surveys conducted in the North‐East Atlantic Ocean during winter and summer 2012, habitats selected by immatures and adult birds were modelled independently, linking gannet density to a set of oceanographic and physiographic predictors. Their large‐scale seasonal distribution was then predicted. We found that gannets displayed a strong season‐dependent competition between immatures and adults, as a consequence of immatures and adults using similar habitats in both summer and winter. During summer, when adults are constrained by reproduction, both groups were spatially highly segregated despite similar habitat preferences (thermal fronts), with youngest individuals selecting habitats out of range of central‐place foragers, highlighting intra‐specific competition. Contrastingly during winter, when reproductive constraints disappear, immature and adult distributions largely overlapped. Our study provides new insights into the role played by age, foraging experience and reproductive constraints on the distribution of marine predators. More specifically, these results highlight in seabirds how the youngest fraction mitigates, through spatial segregation, the competition with experienced adults, and suggest a progressive strategy along the maturation process.     

Trophic segregation of Falkland Islands seabirds: insights from stable isotope analysis

Seabird colonies provide rare opportunities to study trophic segregation in an entire bird community. We here present data on nitrogen and carbon isotope ratios of eight species of seabirds from New Island, Falkland Islands, and compare trophic levels (TL) and foraging distributions. We included adult feathers representing the interbreeding season, as well as chick feathers or down representing the breeding season. The stable isotope ratios indicated differences in feeding areas and TLs between species, consistent with the data of previous conventional diet analyses and observations at sea. We further reviewed conventional and stable isotope seabird community studies calculating the means and ranges of TLs observed across these studies. The mean TL (3.7) of the seabird community on New Island was at the lower end of the mean value range (3.5–4.5), but not significantly different, from the reviewed seabird communities. Seabirds on New Island had a range of 1.3 TLs, which is on the upper end of ranges within a community (0.4–1.5), indicating strong trophic structuring.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

GPS tracking of the foraging movements of Manx Shearwaters Puffinus puffinus breeding on Skomer Island, Wales

We report the first successful use of miniature Global Positioning System loggers to track the ocean‐going behaviour of a c. 400 g seabird, the Manx Shearwater Puffinus puffinus. Breeding birds were tracked over three field seasons during the incubation and chick‐rearing periods on their foraging excursions from the large colony on Skomer Island, Pembrokeshire, UK. Foraging effort was concentrated in the Irish Sea. Likely foraging areas were identified to the north, and more diffusely to the west of the colony. No foraging excursions were recorded significantly to the south of the colony, conflicting with the conclusions of earlier studies based on ringing recoveries and observations. We discuss several explanations including the hypothesis that foraging may have shifted substantially northwards in recent decades. We found no obvious relationship between birds’ positions and water depth, although there was a suggestion that observations at night were in shallower water than those during the day. We also found that, despite the fact that Shearwaters can be observed rafting off‐shore from their colonies in the hours prior to making landfall at night, breeding birds are usually located much further from the colony in the last 8 h before arrival, a finding that has significance for the likely effectiveness of marine protection areas if they are only local to the colony. Short sequences of precise second‐by‐second fixes showed that movement speeds were bimodal, corresponding to sitting on the water (most common at night and around midday) and flying (most common in the morning and evening), with flight behaviour separable into erratic (indicative of searching for food) and directional (indicative of travelling). We also provide a first direct measurement of mean flight speed during directional flight (c. 40 km/h), slower than a Shearwater's predicted maximum range velocity, suggesting that birds are exploiting wave or dynamic soaring during long‐distance travel.