Individual specialization in diet by a generalist marine predator reflects specialization in foraging behaviour

1. We studied chick diet in a known-age, sexed population of a long-lived seabird, the Brünnich's guillemot (Uria lomvia), over 15 years (N = 136; 1993-2007) and attached time-depth-temperature recorders to examine foraging behaviour in multiple years (N = 36; 2004-07). 2. Adults showed specialization in prey fed to offspring, described by multiple indices calculated over 15 years: 27% of diet diversity was attributable to among-individual variation (within-individual component of total niche width = 0.73); average similarity of an individual's diet to the overall diet was 65% (mean proportional similarity between individuals and population = 0.65); diet was significantly more specialized than expected for 70% of individuals (mean likelihood = 0.53). These indices suggest higher specialization than the average for an across-taxa comparison of 49 taxa. 3. Foraging behaviour varied along three axes: flight time, dive depth and dive shape. Individuals showed specialized individual foraging behaviour along each axis. These foraging strategies were reflected in the prey type delivered to their offspring and were maintained over scales of hours to years. 4. Specialization in foraging behaviour and diet was greater over short time spans (hours, days) than over long time spans (years). Regardless of sex or age, the main component of variation in foraging behaviour and chick diet was between individuals. 5. Plasma stable isotope values were similar across years, within a given individual, and variance was low relative to that expected from prey isotope values, suggesting adult diet specialized across years. Stable isotope values were similar among individuals that fed their nestlings similar prey items and there was no difference in trophic level between adults and chicks. We suggest that guillemots specialize on a single foraging strategy regardless of whether chick-provisioning and self-feeding. With little individual difference in body mass and physiology, specialization likely represents learning and memorizing optimal feeding locations and behaviours. 6. There was no difference in survival or reproductive success between specialists and generalists, suggesting these are largely equivalent strategies in terms of evolutionary fitness, presumably because different strategies were advantageous at different levels of prey abundance or predictability. The development of individual specialization may be an important precursor to diversification among seabirds.     

Distribution Patterns Predict Individual Specialization in the Diet of Dolphin Gulls

Many animals show some degree of individual specialization in foraging strategies and diet. This has profound ecological and evolutionary implications. For example, populations containing diverse individual foraging strategies will respond in different ways to changes in the environment, thus affecting the capacity of the populations to adapt to environmental changes and to diversify. However, patterns of individual specialization have been examined in few species. Likewise it is usually unknown whether specialization is maintained over time, because examining the temporal scale at which specialization occurs can prove difficult in the field. In the present study, we analyzed individual specialization in foraging in Dolphin Gulls Leucophaeus scoresbii, a scavenger endemic to the southernmost coasts of South America. We used GPS position logging and stable isotope analyses (SIA) to investigate individual specialization in feeding strategies and their persistence over time. The analysis of GPS data indicated two major foraging strategies in Dolphin Gulls from New I. (Falkland Is./Islas Malvinas). Tagged individuals repeatedly attended either a site with mussel beds or seabird and seal colonies during 5 to 7 days of tracking. Females foraging at mussel beds were heavier than those foraging at seabird colonies. Nitrogen isotope ratios (δ¹⁵N) of Dolphin Gull blood cells clustered in two groups, showing that individuals were consistent in their preferred foraging strategies over a period of at least several weeks. The results of the SIA as well as the foraging patterns recorded revealed a high degree of specialization for particular feeding sites and diets by individual Dolphin Gulls. Individual differences in foraging behavior were not related to sex. Specialization in Dolphin Gulls may be favored by the advantages of learning and memorizing optimal feeding locations and behaviors. Specialized individuals may reduce search and handling time and thus, optimize their energy gain and/or minimize time spent foraging.     

Seasonal Variation in Parental Care Drives Sex-Specific Foraging by a Monomorphic Seabird

Evidence of sex-specific foraging in monomorphic seabirds is increasing though the underlying mechanisms remain poorly understood. We investigate differential parental care as a mechanism for sex-specific foraging in monomorphic Common Murres (Uria aalge), where the male parent alone provisions the chick after colony departure. Using a combination of geolocation-immersion loggers and stable isotopes, we assess two hypotheses: the reproductive role specialization hypothesis and the energetic constraint hypothesis. We compare the foraging behavior of females (n = 15) and males (n = 9) during bi-parental at the colony, post-fledging male-only parental care and winter when parental care is absent. As predicted by the reproductive role specialization hypothesis, we found evidence of sex-specific foraging during post-fledging only, the stage with the greatest divergence in parental care roles. Single-parenting males spent almost twice as much time diving per day and foraged at lower quality prey patches relative to independent females. This implies a potential energetic constraint for males during the estimated 62.8 ± 8.9 days of offspring dependence at sea. Contrary to the predictions of the energetic constraint hypothesis, we found no evidence of sex-specific foraging during biparental care, suggesting that male parents did not forage for their own benefit before colony departure in anticipation of post-fledging energy constraints. We hypothesize that unpredictable prey conditions at Newfoundland colonies in recent years may limit male parental ability to allocate additional time and energy to self-feeding during biparental care, without compromising chick survival. Our findings support differential parental care as a mechanism for sex-specific foraging in monomorphic murres, and highlight the need to consider ecological context in the interpretation of sex-specific foraging behavior.     

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.     

Examining the link between dietary specialization and foraging modes of stream‐dwelling brown trout Salmo trutta

The aim of this study was to explore differences in dietary specialization across two foraging modes (benthic v. surface‐drift foraging) of stream‐dwelling brown trout Salmo trutta. The degree of inter‐individual niche variation within each foraging mode was high, but the dietary specialization was maintained between foraging modes. This study supports the view that if aquatic invertebrates are more abundant and accessible than surface prey, the individuals will not specialize on surface prey (surface‐drift foraging).     

Interpopulation resource partitioning of Lesser Frigatebirds and the influence of environmental context

Conspecific individuals inhabiting nearby breeding colonies are expected to compete strongly for food resources owing to the constraints imposed by shared morphology, physiology, and behavior on foraging strategy. Consequently, colony‐specific foraging patterns that effectively partition the available resources may be displayed. This study aimed to determine whether intraspecific resource partitioning occurs in two nearby colonies of Lesser Frigatebirds (Fregata ariel). A combination of stable isotope analysis and GPS tracking was used to assess dietary and spatial partitioning of foraging resources during the 2013 and 2014 breeding seasons. These results were compared to vessel‐derived estimates of prey availability, local primary productivity, and estimates of reproductive output to suggest potential drivers and implications of any observed partitioning. Isotopic data indicated a more neritic source of provisioned resources for near‐fledged chicks at an inshore colony, whereas their offshore counterparts were provisioned with resources with a more pelagic signal. Deep pelagic waters (>200 m) had higher availability of a preferred prey type despite a trend for lower primary productivity. Differences in foraging ecology between the two populations may have contributed to markedly different reproductive outputs. These findings suggest environmental context influences dietary and spatial aspects of foraging ecology. Furthermore, the effect of colony‐specific foraging patterns on population demography warrants further research.     

Consequences of foraging trip duration on provisioning behaviour and fledging condition of common murres Uria aalgae

We examine the provisioning constraints of a pursuit‐diving seabird in a cold ocean regime by comparing the behaviour of common murres Uria aalge rearing chicks at two colonies in the Northwest Atlantic during 1998‐2000. Funk Island is the largest (340,000–400,000 breeding pairs) and most offshore (60 km) colony of common murres in eastern Canada. Seventy‐five percent of the Northwest Atlantic population of common murres breeds on this island. Great Island is one island within the Witless Bay Ecological Reserve, which is the second largest breeding aggregation (100 000 breeding pairs) and is located near‐shore (2 km). The primary forage fish species in Newfoundland waters is capelin Mallotus villosus, which spawns on or near coastal beaches during summer. Therefore, the two study colonies differ in their distance to food resources and colony size. It is within this natural context that we compare: (1) prey types and frequency of delivery (amount of prey), (2) parental time budgets, and (3) the mass and condition (mass/wing length) of fledglings at both colonies. Similarly sized female capelin (100–150 mm) were delivered to chicks at both colonies. Foraging time per day per parent, a proxy of foraging effort, was similar at both colonies (Great Island: 5.1 h; Funk Island: 5.5 h), as was the percentage of time spent with mates (Great Island: 12.3%; Funk Island: 10.9%). Foraging trips, however, were longer at Funk Island (4.1 h) than at Great Island (2.9 h). This resulted in lower feeding rates of chicks (0.17 feeds per h) and poorer condition of fledglings (2.9 g/mm) at Funk Island compared to those at Great Island (0.22 feeds per h; 3.9 g/mm). We hypothesize that provisioning efforts are constrained at Funk Island by (1) distant food resources and increased competitor density, resulting in longer foraging trip durations and (2) the time spent paired with mates at the colony, which may reflect a minimum time required to maintain breeding sites due to higher breeding densities at Funk Island compared to Great Island. Demographic consequences of this poor fledgling condition at Funk Island are unknown, but fledglings may sufficiently accelerate growth at sea due to their closer proximity to an important nursery area. If fledgling survival is compromised, however, the lower potential for growth at Funk Island will impact the entire Northwest Atlantic population of common murres.     

The foraging decisions of a central place foraging seabird in response to fluctuations in local prey conditions

During reproduction, seabirds need to balance the demands of self- and offspring-provisioning within the constraints imposed by central place foraging. To assess behavioral adjustments and tolerances to these constraints, we studied the feeding tactics and reproductive success of common murres (also known as common guillemots) Uria aalge , at their largest and most offshore colony (Funk Island) where parents travel long distances to deliver a single capelin Mallotus villosus to their chicks. We assessed changes in the distance murres traveled from the colony, their proximate foraging locations and prey size choice during two successive years in which capelin exhibited an order of magnitude decrease in density and a shift from aggregated (2004) to dispersed (2005) distributions. When capelin availability was low (2005), parental murres increased their maximum foraging distances by 35% (60 to 81 km) and delivered significantly larger capelin to chicks, as predicted by central place foraging theory. Murres preferred large (>140 mm) relative to small capelin (100–140 mm) in both years, but unexpectedly this preference increased as the relative density of large capelin decreased. We conclude that single prey-loading murres target larger capelin during long foraging trips as parents are 'forced' to select the best prey for their offspring. Low fledgling masses suggest also that increased foraging time when capelin is scarce may come at a cost to the chicks (i.e. fewer meals per day). Murres at this colony may be functioning near physiological limits above which further or sustained adjustments in foraging effort could compromise the life-time reproductive success of this long-lived seabird.     

Metapopulation dynamics and foraging plasticity in a highly vagile seabird,the southern rockhopper penguin

Population connectivity is driven by individual dispersal potential and modulated by natal philopatry. In seabirds, high vagility facilitates dispersal yet philopatry is also common, with foraging area overlap often correlated with population connectivity. We assess the interplay between these processes by studying past and current connectivity and foraging niche overlap among southern rockhopper penguin colonies of the coast of southern South America using genomic and stable isotope analyses. We found two distinct genetic clusters and detected low admixture between northern and southern colonies. Stable isotope analysis indicated niche variability between colonies, with Malvinas/Falklands colonies encompassing the species entire isotopic foraging niche, while the remaining colonies had smaller, nonoverlapping niches. A recently founded colony in continental Patagonia differed in isotopic niche width and position with Malvinas/Falklands colonies, its genetically identified founder population, suggesting the exploitation of novel foraging areas and/or prey items. Additionally, dispersing individuals found dead across the Patagonian shore in an unusual mortality event were also assigned to the northern cluster, suggesting northern individuals reach southern localities, but do not breed in these colonies. Facilitated by variability in foraging strategies, and especially during unfavorable conditions, the number of dispersing individuals may increase and enhance the probability of founding new colonies. Metapopulation demographic dynamics in seabirds should account for interannual variability in dispersal behavior and pay special attention to extreme climatic events, classically related to negative effects on population trends.     

Specialized foraging habits of adult female California sea lions Zalophus californianus

Individual foraging specialization occurs when organisms use a small subset of the resources available to a population. This plays an important role in population dynamics since individuals may have different ecological functions within an ecosystem related to habitat use and prey preferences. The foraging habitat fidelity and degree of specialization of California sea lions (Zalophus californianus) were evaluated by analyzing the stable isotopes values of carbon and nitrogen in vibrissae collected from 16 adult females from the reproductive colony on Santa Margarita Island, Magdalena Bay, Mexico, in 2012 and 2013. Based on the degree of individual specialization in δ¹⁵N, 62.5% of the females assessed can be considered specialist consumers focusing on the same prey or different prey from the same trophic level. The degree of individual specialization in δ¹³C indicated that 100% of the individuals showed fidelity to their foraging habitat as some fed in the lagoon, others foraged along the coast, and a third group preferred prey from the pelagic environment during both the breeding and nonbreeding seasons, suggesting diversification of foraging areas. Foraging area fidelity persisted despite the 2°C increase in the sea surface temperature over the course of the study period.     

Mapping pre- and post- fledging foraging locations of thick-billed murres in the North Water polynya

In this study of thick- billed murres in high- arctic Greenland we used electronic data loggers and satellite transmitters (PTTs) to identify the foraging areas of chick-rearing adults, and to map the routes and staging areas of adults accompanying post-fledging chicks during their swimming migration within the North Water (NOW) polynya. During the pre-fledging period the majority of 19 foraging trips performed by 8 birds went to a shelf area north of the colony where 83% of all dives took place. Individual birds headed in different directions during successive trips, and went up to 47 km from the colony. Upon fledging the four PTT-tagged adult/chick pairs initiated swimming migration by heading south-west from the colony. All pairs moved fast until they arrived at a shallow bank area ca 180 km from the colony, where at least two of the pairs remained for more than a week. Speed during the active migration averaged 2.5 km h^-1 with a peak of 6.6 km h^-1. In the pre-fledging period the birds utilised a feeding area outside the normal foraging range of murres from other colonies. Similarly, post-fledging adult/chick pairs may have benefitted from reduced food competition when they moved to a staging area situated at the only shallow area in the polynya without any adjacent murre colony. This initial study suggested that the high-arctic murres did not hasten towards the wintering grounds, and that the NOW remained important even to post-fledging murres.     

Foraging ecology of the endangered wood stork recorded in the stable isotope signature of feathers

Down feathers and regurgitant were collected from nestling wood storks (Mycteria americana) from two inland and two coastal breeding colonies in Georgia. The stable isotopic ratios of carbon (¹³C/¹²C) and nitrogen (¹⁵N/¹⁴N) in these materials were analyzed to gain insights into the natal origins of juvenile storks and the foraging activities of adults. Down feathers differed in '¹³C between inland and coastal colonies, having average isotopic values that reflected the sources of carbon fixed in biomass at the base of the food web. Feathers from the inland colonies differed between colonies in '¹⁵N, while those from the coastal colonies did not. These patterns primarily reflected the foraging activities of parent storks, with individuals capturing differing percentages of prey of distinct trophic status at each colony. Collectively, the carbon and nitrogen isotopic signatures of feather keratin were used to distinguish nestlings from each colony, except for instances where storks from different colonies foraged in common wetlands. The stable isotopic composition of food items in regurgitant was used to reconstruct the trophic structure of the ecosystems in which wood storks foraged. Predicted foraging activities based on the isotopic composition of keratin were generally consistent with the percentage of prey types (freshwater vs. saltwater and lower trophic level vs. upper trophic level consumer) observed in regurgitant, except for the coastal colony at St. Simons Island, where the '¹³C of feathers strongly suggested that freshwater prey were a significant component of the diet. This inconsistency was resolved by aerial tracking of adults during foraging excursions using a fixed-wing aircraft. Observed foraging activities supported interpretations based on the stable isotope content of feathers, suggesting that the latter provided a better record of overall foraging activity than regurgitant analysis alone. Observed foraging patterns were compared to the predictions of a statistical model that determined habitat utilization based on habitat availability using a geographic information system (GIS) database. Observed foraging activities and those predicted from feathers both suggested that some adult storks preferred to feed their young freshwater prey, even when saltwater resources were more accessible in the local environment. This conclusion supports the contention that wood stork populations are sensitive to changes in the distribution of freshwater habitats along the southeastern coastal plain of the United States.     

Variation in trophic resources in female South American sea lions at a small geographic scale

Difference among colonies in the population structure of otariids can be driven by philopatry and/or by specializations in the foraging ecology of females. In northern Patagonia, the South American sea lion (SASL) shows some degree of spatial genetic structure among colonies from north and south zones. This study aims to explore the isotopic niche of SASL females in the last period of the pregnancy from different colonies of northern Patagonia and to consider whether the fine scale genetic spatial structuring is potentially related to variation in trophic resources. Stable isotope analysis was performed on 101 skin samples of newborn pups in 10 colonies, as a proxy for the feeding ecology of their mothers. Differences among colonies in the metrics studied revealed the plasticity of the species and support individual trophic specialization of SASL females at a small geographic scale. Also, significant differences were found in all isotopic metrics between the north and south zones. Several hypotheses were proposed to explain the differences in SASL females' isotope values (e.g., use of different foraging areas or prey, isotopic baseline variation). Nonetheless, further research is needed to better understand the relation between fine scale genetic structuring and the foraging ecology of SASL females.     

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.     

Within-colony feeding selectivity by a corallivorous reef fish: foraging to maximize reward?

Foraging theory predicts that individuals should choose a prey that maximizes energy rewards relative to the energy expended to access, capture, and consume the prey. However, the relative roles of differences in the nutritive value of foods and costs associated with differences in prey accessibility are not always clear. Coral‐feeding fishes are known to be highly selective feeders on particular coral genera or species and even different parts of individual coral colonies. The absence of strong correlations between the nutritional value of corals and prey preferences suggests other factors such as polyp accessibility may be important. Here, we investigated within‐colony feeding selectivity by the corallivorous filefish, Oxymonacanthus longirostris, and if prey accessibility determines foraging patterns. After confirming that this fish primarily feeds on coral polyps, we examined whether fish show a preference for different parts of a common branching coral, Acropora nobilis, both in the field and in the laboratory experiments with simulated corals. We then experimentally tested whether nonuniform patterns of feeding on preferred coral species reflect structural differences between polyps. We found that O. longirostris exhibits nonuniform patterns of foraging in the field, selectively feeding midway along branches. On simulated corals, fish replicated this pattern when food accessibility was equal along the branch. However, when food access varied, fish consistently modified their foraging behavior, preferring to feed where food was most accessible. When foraging patterns were compared with coral morphology, fish preferred larger polyps and less skeletal protection. Our results highlight that patterns of interspecific and intraspecific selectivity can reflect coral morphology, with fish preferring corals or parts of coral colonies with structural characteristics that increase prey accessibility.     

Divergent diving behavior during short and long trips of a bimodal forager,the little auk Alle alle

The purpose of this study was to characterize for the first time seabird diving behavior during bimodal foraging. Little auks Alle alle, small zooplanktivorous Alcids of the High Arctic, have recently been shown to make foraging trips of short and long duration. Because short (ST) and long trips (LT) are thought to occur in different locations and serve different purposes (chick‐ and self‐feeding, respectively) we hypothesized that foraging differences would be apparent, both in terms of water temperature and diving characteristics. Using Time Depth Recorders (TDRs), we tested this hypothesis at three colonies along the Greenland Sea with contrasting oceanographic conditions. We found that diving behavior generally differed between ST and LT. However, the magnitude of the disparity in diving characteristics depended on local foraging conditions. At the study site where conditions were favorable, diving behavior differed only to a small degree between LT and ST. Together with a lack of difference in diving depth and ocean temperature, this indicates that these birds did not increase their foraging effort during ST nor did they travel long distances to seek out more profitable prey. In contrast, where local foraging conditions were poor, birds increased their diving effort substantially to collect a chick meal during ST as indicated by longer, more U‐shaped dives with slower ascent rates and shorter resting times (post‐dive intervals and extended surface pauses). In addition, large differences in diving depth and ocean temperature indicate that birds forage on different prey species and utilize different foraging areas during LT, which may be up to 200 km away from the colony. Continued warming and deteriorating near‐colony foraging conditions may have energetic consequences for little auks breeding in the eastern Greenland Sea.     

Is There Division of Labor in Cooperative Pseudoscorpions? An Analysis of the Behavioral Repertoire of a Tropical Species

Division of labor is a strategy that maximizes the foraging and reproductive success of eusocial insects. Although some arachnids exhibit colony structure and social organization similar to that of hymenopterans, temporal polyethism has only been demonstrated in few species. The social organization of cooperative pseudoscorpions Paratemnoides nidificator is similar to that of social spiders, but it involves a clear division of labor. Work allocation was experimentally investigated in colonies composed of only one developmental stage (young or adults) or by one sex (males or females), through laboratory manipulation. During 44 h of observation, more than 14 000 behavioral repetitions were quantified, distributed in 95 different types of behavioral acts, and grouped in 10 behavioral categories. The results showed that reproductive colonies of P. nidificator are maintained by gender‐ and age‐based activities. Males and non‐reproductive females performed the external cleaning of the colony and prey capture. Reproductive females take care of the juveniles and build reproductive silk chambers. Nymphs build most of the molt chambers and perform internal cleaning. In the absence of nymphs, male colonies survived 1–2 mo, while female colonies survived 3–4 mo. In nymph colonies, work is readjusted so that all maintenance tasks are executed. This is the first study clearly demonstrating division of tasks in arachnids. It suggests that specialization is an adaptative and evolutionarily old trait in this species. Unlike cooperative spiders, P. nidificator possesses physiological (e.g. reproduction, ecdysis, lifespan) and behavioral (e.g. behavioral synchrony or self‐organization) characteristics that allow task specialization.     

Prey capture and selection throughout the breeding season in a deep‐diving generalist seabird,the thick‐billed murre

Generalist seabirds forage on a variety of prey items providing the opportunity to monitor diverse aquatic fauna simultaneously. For example, the coupling of prey consumption rates and movement patterns of generalist seabirds might be used to create three‐dimensional prey distribution maps (‘preyscapes’) for multiple prey species in the same region. However, the complex interaction between generalist seabird foraging behaviour and the various prey types clouds the interpretation of such preyscapes, and the mechanisms underlying prey selection need to be understood before such an application can be realized. Central place foraging theory provides a theoretical model for understanding such selectivity by predicting that larger prey items should be 1) selected farther from the colony and 2) for chick‐feeding compared with self‐feeding, but these predictions remain untested on most seabird species. Furthermore, rarely do we know how foraging features such as handling time, capture methods or choice of foraging location varies among prey types. We used three types of animal‐borne biologgers (camera loggers, GPS and depth‐loggers) to examine how a generalist Arctic seabird, the thick‐billed murre Uria lomvia, selects and captures their prey throughout the breeding season. Murres captured small prey at all phases of a dive, including while descending and ascending, but captured large fish mostly while ascending, with considerably longer handling times. Birds captured larger prey and dove deeper during chick‐rearing. As central place foraging theory predicted, birds travelling further also brought bigger prey items for their chick. The location of a dive (distance from colony and distance to shore) best explained which prey type was the most likely to get caught in a dive, and we created a preyscape surrounding our study colony. We discuss how these findings might aid the use of generalist seabirds as bioindicators.     

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.     

Large‐scale molecular barcoding of prey DNA reveals predictors of intrapopulation feeding diversity in a marine predator

Predator–prey interactions are critical in understanding how communities function. However, we need to describe intraspecific variation in diet to accurately depict those interactions. Harbor seals (Phoca vitulina) are an abundant marine predator that prey on species of conservation concern. We estimated intrapopulation feeding diversity (variation in feeding habits between individuals of the same species) of harbor seals in the Salish Sea. Estimates of feeding diversity were examined relative to sex, month, and location using a novel approach that combined molecular techniques, repeated cross‐sectional sampling of scat, and a specialization metric (within‐individual consistency in diet measured by the Proportional Similarity Index ()). Based on 1,083 scat samples collected from five haul‐out sites during four nonsequential years, we quantified diet using metabarcoding techniques and determined the sex of the scat depositor using a molecular assay. Results suggest that intrapopulation feeding diversity was present. Specialization was high over short periods (24–48 hr,  = 0.392, 95% CI = 0.013, R = 100,000) and variable in time and space. Females showed more specialization than males, particularly during summer and fall. Additionally, demersal and benthic prey species were correlated with more specialized diets. The latter finding suggests that this type of prey likely requires specific foraging strategies and that there are trade‐offs between pelagic and benthic foraging styles for harbor seals. This differential feeding on prey species, as well as between sexes of harbor seals, indicates that predator–prey interactions in harbor seals are complex and that each sex may have a different impact on species of conservation concern. As such, describing intrapopulation feeding diversity may unravel hitherto unknown complex predator–prey interactions in the community.