Foraging range scales with colony size in high-latitude seabirds.

Density-dependent prey depletion around breeding colonies has long been considered an important factor controlling the population dynamics of colonial animals.1-4 Ashmole proposed that as seabird colony size increases, intraspecific competition leads to declines in reproductive success, as breeding adults must spend more time and energy to find prey farther from the colony.1 Seabird colony size often varies over several orders of magnitude within the same species and can include millions of individuals per colony.5,6 As such, colony size likely plays an important role in determining the individual behavior of its members and how the colony interacts with the surrounding environment.6 Using tracking data from murres (Uria spp.), the world's most densely breeding seabirds, we show that the distribution of foraging-trip distances scales to colony size0.33 during the chick-rearing stage, consistent with Ashmole's halo theory.1,2 This pattern occurred across colonies varying in size over three orders of magnitude and distributed throughout the North Atlantic region. The strong relationship between colony size and foraging range means that the foraging areas of some colonial species can be estimated from colony sizes, which is more practical to measure over a large geographic scale. Two-thirds of the North Atlantic murre population breed at the 16 largest colonies; by extrapolating the predicted foraging ranges to sites without tracking data, we show that only two of these large colonies have significant coverage as marine protected areas. Our results are an important example of how theoretical models, in this case, Ashmole's version of central-place-foraging theory, can be applied to inform conservation and management in colonial breeding species.

Different individual-level responses of great black-backed gulls (Larus marinus) to shifting local prey availability.

To grow, survive and reproduce under anthropogenic-induced changes, individuals must respond quickly and favourably to the surrounding environment. A species that feeds on a wide variety of prey types (i.e. generalist diet) may be comprised of generalist individuals, specialist individuals that feed on different prey types, or a combination of the two. If individuals within a population respond differently to an environmental change, population-level responses may not be detectable. By tracking foraging movements of great black-backed gulls (Larus marinus), a generalist species, we compared group-level and individual-level responses to an increase in prey biomass (capelin; Mallotus villosus) during the breeding season in coastal Newfoundland, Canada. As hypothesized, shifts in prey availability resulted in significantly different individual responses in foraging behaviour and space use, which was not detectable when data from individuals were combined. Some individuals maintained similar foraging areas, foraging trip characteristics (e.g., trip length, duration) and habitat use with increased capelin availability, while others shifted foraging areas and habitats resulting in either increased or decreased trip characteristics. We show that individual specialization can be non-contextual in some gulls, whereby these individuals continuously use the same feeding strategy despite significant change in prey availability conditions. Findings also indicate high response diversity among individuals to shifting prey conditions that a population- or group-level study would not have detected, emphasizing the importance of examining individual-level strategies for future diet and foraging studies on generalist species.

Allopatric humpback whales of differing generations share call types between foraging and wintering grounds.

Humpback whales (Megaptera novaeangliae) are a cosmopolitan baleen whale species with geographically isolated lineages. Despite last sharing an ancestor ~ 2-3 million years ago, Atlantic and Pacific foraging populations share five call types. Whether these call types are also shared between allopatric breeding and foraging populations is unclear, but would provide further evidence that some call types are ubiquitous and fixed. We investigated whether these five call types were present on a contemporary foraging ground (Newfoundland, 2015-2016) and a historic breeding ground (Hawaii, 1981-1982). Calls were classified using aural/visual (AV) characteristics; 16 relevant acoustic variables were measured and a Principal Component Analysis (PCA) was used to examine within-call and between-population variation. To assess whether between-population variation influenced classification, all 16 variables were included in classification and regression tree (CART) and random forest analyses (RF). All five call types were identified in both populations. Between-population variation in combined acoustic variables (PC1, PC2, PC3) was lower within call types than among call types, and high agreement between AV and quantitative classification (CART: 83% agreement; RF: 77% agreement) suggested that acoustic characteristics were more similar within than among call types. Findings indicate that these five call types are shared across allopatric populations, generations, and behavioural contexts.

Isotopic Discrimination (δ15N, δ13C) in Captive and Wild Common Murres (Uria aalge) and Atlantic Puffins (Fratercula arctica).

Studying the diet of consumers using stable isotopes provides insight into the foraging ecology of individuals and species. To accurately reconstruct the integrated diet of animals using stable isotope values, we must quantify diet-tissue discrimination factors (DTDFs), or the way in which stable isotopes in prey are incorporated into the tissues of consumers. To quantify DTDFs, controlled experiments are needed, whereby consumers are fed a constant diet. However, relatively few controlled-diet studies have been conducted for seabirds. In this study, captive adult Atlantic puffins (Fratercula arctica) and common murres (Uria aalge) were fed a two-source diet of capelin (Mallotus villosus) and Atlantic silverside (Menidia menidia) to determine the DTDFs for the cellular component of blood and plasma for both δ15N and δ13C. The DTDFs for the cellular component (Δ15N: 2.80±0.28; Δ13C: 1.21±0.22) and plasma (Δ15N: 1.72±1.03; Δ13C: -0.18±0.56) of puffins were similar to those for the cellular component (Δ15N: 2.91±0.18; Δ13C: 1.09±0.23) and plasma (Δ15N: 2.18±0.77; Δ13C: -0.70±0.18) of murres. We reconstructed the diet of wild murres and puffins breeding on the northeastern coast of Newfoundland using previously published DTDFs and estimated DTDFs from our feeding experiment. Reconstructed dietary proportions supported a priori knowledge of diet, although outputs were sensitive to the DTDF used. Despite the similarity of our DTDFs for puffins and murres, along with the similarity of our DTDFs with those of other seabird species, our sensitivity analysis revealed considerable differences among resultant dietary contributions from mixing models, further highlighting the importance of using species- and tissue-specific DTDFs to enhance knowledge in the foraging ecology of seabirds using stable isotopes.

Diet-tissue discrimination factors (δ15 N and δ13 C values) for blood components in Magellanic (Spheniscus magellanicus) and southern rockhopper (Eudyptes chrysocome) penguins.

RATIONALE: Analysis of the stable isotope ratios of carbon and nitrogen (δ13 C and δ15 N values) is increasingly being used to gain insight into predator trophic ecology, which requires accurate diet-tissue discrimination factors (DTDFs), or the isotopic difference between prey and predator. Accurate DTDFs must be calculated from predators consuming an isotopically constant diet over time in controlled feeding experiments, but these studies have received little attention to date, especially among seabird species. METHODS: In this study, aquarium-housed Magellanic (Spheniscus magellanicus) and southern rockhopper (Eudyptes chrysocome) penguins were fed a single-prey source diet (capelin Mallotus villosus) for eight weeks. Stable isotope ratios (δ13 C and δ15 N values) of penguin blood (cellular component and plasma) and capelin were measured using mass spectrometry and then used to calculate DTDFs for both components of penguin blood by comparison with prey values. RESULTS: The DTDFs for plasma were -0.63 ± 0.49 (mean ± SD) and -0.27 ± 0.22 for δ13 C values, and 2.60 ± 0.50 and 2.78 ± 0.22 for δ15 N values for Magellanic and southern rockhopper penguins, respectively, while the DTDFs for the cellular component were 1.22 ± 0.03 and 1.26 ± 0.03 for δ13 C values, and 2.54 ± 0.07 and 2.43 ± 0.17 for δ15 N values. CONCLUSIONS: We compare our DTDFs with published values from blood components of penguins and discuss the effects that lipid extraction, sample storage, and diet have on the DTDFs of penguin blood components. This study provides accurate DTDFs of blood components for two seabird species of conservation concern, and is one of the first to provide plasma DTDFs for penguins, which are underrepresented in the seabird literature.

Egg cannibalism in capelin Mallotus villosus at beach and deep-water spawning habitats in the north-west Atlantic Ocean.

We investigated egg cannibalism in spawning capelin Mallotus villosus on the north-east Newfoundland coast during July 2012-2014, specifically whether sex, spawning condition (i.e., spawning or spent) and spawning habitat influenced egg cannibalism. Capelin spawning in deep-water were 4.5-14 times more likely to be cannibals than those at the beach, probably due to the higher spatial overlap of spawners and conspecific eggs within this habitat relative to beaches. Males were 2.1-3.7 times more likely to be cannibals than females, but female cannibals had more eggs per stomach. Spawning fish were 1.6-1.9 times more likely to be cannibals than spent fish, but spent female cannibals had more eggs per stomach relative to spawning males and females in either habitat. Findings suggest that cannibalism may be an important foraging strategy, especially at deep-water spawning habitat, possibly extending the spawning season for males or increasing the probability of post-spawning survival for females. Although 44% of sampled females and 50% of males were cannibals, the estimated mortality due to egg cannibalism was low (0.49-2.97% of eggs produced annually), suggesting that egg cannibalism does not influence recruitment to a great extent in Newfoundland capelin.

High flight costs, but low dive costs, in auks support the biomechanical hypothesis for flightlessness in penguins.

Flight is a key adaptive trait. Despite its advantages, flight has been lost in several groups of birds, notably among seabirds, where flightlessness has evolved independently in at least five lineages. One hypothesis for the loss of flight among seabirds is that animals moving between different media face tradeoffs between maximizing function in one medium relative to the other. In particular, biomechanical models of energy costs during flying and diving suggest that a wing designed for optimal diving performance should lead to enormous energy costs when flying in air. Costs of flying and diving have been measured in free-living animals that use their wings to fly or to propel their dives, but not both. Animals that both fly and dive might approach the functional boundary between flight and nonflight. We show that flight costs for thick-billed murres (Uria lomvia), which are wing-propelled divers, and pelagic cormorants (Phalacrocorax pelagicus) (foot-propelled divers), are the highest recorded for vertebrates. Dive costs are high for cormorants and low for murres, but the latter are still higher than for flightless wing-propelled diving birds (penguins). For murres, flight costs were higher than predicted from biomechanical modeling, and the oxygen consumption rate during dives decreased with depth at a faster rate than estimated biomechanical costs. These results strongly support the hypothesis that function constrains form in diving birds, and that optimizing wing shape and form for wing-propelled diving leads to such high flight costs that flying ceases to be an option in larger wing-propelled diving seabirds, including penguins.

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.

Effects of gill-net fishing on marine birds in a biological hotspot in the northwest Atlantic.

Marine biological hotspots, or areas where high abundances of species overlap in space and time, are ecologically important areas because energy flow through marine food webs, a key ecosystem process, is maximized in these areas. I investigated whether top predators aggregated at persistent spawning sites of a key forage fish species, capelin (Mallotus villosus), on the NE coast of Newfoundland during July and August 2000-2003. By examining the distributional patterns of top predators through ship-based surveys at multiple spatial and temporal scales, I found that the biomasses of birds-dominated by Common Murres (Uria aalge)-and mammals-dominated by whale species-were concentrated along the coast, with a biological hotspot forming near two persistent spawning sites of capelin in all years. The formation of this hotspot was well defined in space and time from middle of July to middle of August, likely coinciding with the spawning chronology of capelin. Within this hotspot, there was a high spatial and temporal overlap of Common Murres and gill nets set to capture Atlantic cod (Gadus morhua). This resulted in breeding murres becoming entangled in gill nets while feeding on spawning capelin. Despite an acknowledged uncertainty of bycatch mortality, estimates for the larger regional-scale area (1936-4973 murres/year; 0.2-0.6% of the breeding population) underestimated mortality relative to estimates within the hotspot (3053-14054 murres/year; 0.4-1.7%). Although fishing effort for Atlantic cod has declined substantially since the groundfish moratorium in 1992, chronic, unnatural, and additive mortality through bycatch continues in coastal Newfoundland. Restricted use of gill nets within this and other biological hotspots during the capelin spawning period appears to be a straightforward application of the "ecological and biologically significant area" management framework in Canada's Oceans Act. This protection would minimize murre bycatch and maintain ecosystem integrity.