Stable isotopes document seasonal changes in trophic niches and winter foraging individual specialization in diving predators from the Southern Ocean

1. Climatic variation outside the breeding season affects fluctuations in population numbers of seabirds and marine mammals. A challenge in identifying the underlying biological mechanisms is the lack of information on their foraging strategies during winter, when individuals migrate far from their breeding grounds. 2. We investigated the temporal variability in resource partitioning within the guild of five sympatric Subantarctic penguins and fur seals from Crozet Islands. The stable isotopic ratios of carbon (delta(13)C) and nitrogen (delta(15)N) for whole blood were measured for penguins and fur seals, as were the isotopic ratios for penguin nails and food. Animals were sampled at two periods, during breeding in summer and at their arrival in the colonies in spring (hereafter winter, since the temporal integration of blood amounting to several months). 3. In summer, delta(13)C and delta(15)N for blood samples defined three foraging areas and two trophic levels, respectively, characterizing four nonoverlapping trophic niches. King penguins and female Antarctic and Subantarctic fur seals are myctophid eaters foraging in distinct water masses, while both macaroni and rockhopper penguins had identical isotopic signatures indicating feeding on crustaceans near the archipelago. 4. Isotopic ratios were almost identical in summer and winter suggesting no major changes in the species niches, and hence, in the trophic structure of the guild during the nonbreeding period. A seasonal difference, however, was the larger variances in delta(13)C (and also to a lesser extent in delta(15)N) values in winter, thus verifying our hypothesis that trophic niches widen when individuals are no longer central place foragers. 5. Winter isotopic ratios of macaroni penguins and male Antarctic fur seals had large variances, indicating individual foraging specializations. The range of delta(13)C and delta(15)N values of male fur seals showed, respectively, that they dispersed over a wide latitudinal gradient (from Antarctica to north of the archipelago) and fed on different prey (crustaceans and fish). 6. By comparing summer and winter isotopic ratios and examining the summer diet, we highlight the feeding habits of marine predators that were not previously addressed. The findings have a number of implications for understanding the functioning of the pelagic ecosystem and on the demography of these species.     

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.     

Is individual consistency in body mass and reproductive decisions linked to individual specialization in foraging behavior in a long‐lived seabird?

Individual specialization in diet or foraging behavior within apparently generalist populations has been described for many species, especially in polar and temperate marine environments, where resource distribution is relatively predictable. It is unclear, however, whether and how increased environmental variability – and thus reduced predictability of resources – due to global climate change will affect individual specialization. We determined the within‐ and among‐individual components of the trophic niche and the within‐individual repeatability of δ¹³C and δ¹⁵N in feathers and red blood cells of individual female southern rockhopper penguins (Eudyptes chrysocome) across 7 years. We also investigated the effect of environmental variables (Southern Annular Mode, Southern Oscillation Index, and local sea surface temperature anomaly) on the isotopic values, as well as the link between stable isotopes and female body mass, clutch initiation dates, and total clutch mass. We observed consistent red blood cell δ¹³C and δ¹⁵N values within individuals among years, suggesting a moderate degree of within‐individual specialization in C and N during the prebreeding period. However, the total niche width was reduced and individual specialization not present during the premolt period. Despite significant interannual differences in isotope values of C and N and environmental conditions, none of the environmental variables were linked to stable isotope values and thus able to explain phenotypic plasticity. Furthermore, neither the within‐individual nor among‐individual effects of stable isotopes were found to be related to female body mass, clutch initiation date, or total clutch mass. In conclusion, our results emphasize that the degree of specialization within generalist populations can vary over the course of 1 year, even when being consistent within the same season across years. We were unable to confirm that environmental variability counteracts individual specialization in foraging behavior, as phenotypic plasticity in δ¹³C and δ¹⁵N was not linked to any of the environmental variables studied.     

Isotopic discrimination between food and blood and feathers of captive penguins: implications for dietary studies in the wild

Using measurements of naturally occurring stable isotopes to reconstruct diets or source of feeding requires quantifying isotopic discrimination factors or the relationships between isotope ratios in food and in consumer tissues. Diet-tissue discrimination factors of carbon ((13)C/(12)C, or delta (13)C) and nitrogen ((15)N/(14)N, or delta (15)N) isotopes in whole blood and feathers, representing noninvasive sampling techniques, were examined using three species of captive penguins (king Aptenodytes patagonicus, gentoo Pygoscelis papua, and rockhopper Eudyptes chrysocome penguins) fed known diets. King and rockhopper penguins raised on a constant diet of herring and capelin, respectively, had tissues enriched in (15)N compared to fish, with discrimination factors being higher in feathers than in blood. These data, together with previous works, allowed us to calculate average discrimination factors for (15)N between whole lipid-free prey and blood and feathers of piscivorous birds; they amount to +2.7 per thousand and +4.2 per thousand, respectively. Both fish species were segregated by their delta (13)C and delta (15)N values, and importantly, lipid-free fish muscle tissue was consistently depleted in (13)C and enriched in (15)N compared to whole lipid-free fish. This finding has important implications because previous studies usually base dietary reconstructions on muscle of prey rather than on whole prey items consumed by the predator. We tested the effect of these differences using mass balance calculations to the quantification of food sources of gentoo penguins that had a mixed diet. Modeling indicated correct estimates when using the isotopic signature of whole fish (muscle) and the discrimination factors between whole fish (muscle) and penguin blood. Conversely, the use of isotopic signatures of muscle together with discrimination factors between whole fish and blood (or the reverse) leads to spurious estimates in food proportions. Consequently, great care must be taken in the choice of isotopic discrimination factors to apply to wild species for which no controlled experiments on captive individuals have been done. Finally, our results also indicate that there is no need to remove lipids before isotopic analysis of avian blood.     

Seasonal and annual differences in the foraging ecology of two gull species breeding in sympatry and their use of fishery discards

Niche segregation between similar species will result from an avoidance of competition but also from environmental variability, including nowadays anthropogenic activities. Gulls are among the seabirds with greater behavioural plasticity, being highly opportunistic and feeding on a wide range of prey, mostly from anthropogenic origin. Here, we analysed blood and feather stable isotopes combined with pellet analysis to investigate niche partitioning between Audouin's gull Larus audouinii and yellow‐legged gull Larus michahellis breeding in sympatry at Deserta Island, southern Portugal, during 2014 and 2015. During the breeding season there was considerable overlap in the adults’ diet, as their stable isotope values of blood and primary feather (P1) did not differ, and their pellets were comprised mainly by marine fish species. However, Audouin's gulls presented higher occurrences of pelagic fish, while yellow‐legged gulls fed more on demersal fish, insects, and refuse. SIAR mixing models also estimated a higher proportion of demersal fish in the diet of yellow‐legged gulls. We also found differences between the two gull species in chicks’ feathers, suggesting that Audouin's gull adults selected prey with lower carbon isotope values to feed their young. Secondary feather (S8) of Audouin's gull presented higher isotope values compared to yellow‐legged gulls, indicating different foraging areas (δ¹³C) and/ or trophic levels (δ¹⁵N) between the two species in the non‐breeding season. During both the all‐year and non‐breeding periods the yellow‐legged gull showed a broader isotopic niche width than Audouin's gull in 2013, and in 2014 the two gull species exhibited different isotopic niche spaces. Our study suggests that both gull species foraged in association with fisheries during the breeding season. In this sense, a discard ban implemented under the new European Union Common Fisheries Policy may lead to a food shortage, therefore future research should closely monitor the population dynamics of Audouin's and yellow‐legged gulls.     

Breeding Yellow-legged Gulls increase consumption of terrestrial prey after landfill closure

The ecology of opportunistic foragers can be highly dependent on anthropogenic food sources, such as landfills, resulting in changes in several ecological and demographic aspects. The total closure of several landfill sites and the use of deterrence systems to prevent access to the remaining open landfill sites in a region in the northern Iberian Peninsula provided an excellent opportunity to evaluate the consequences of landfills on the trophic ecology of an opportunistic forager, the Yellow-legged Gull Larus michahellis, using these resources. Based on stable isotope analyses, we used mixing models to estimate changes in diet before and after the closure of the majority of landfills in the breeding and the non-breeding season. We found a decrease in the consumption of food from landfills after their closure, which was accompanied by an increase in feeding on terrestrial prey (mostly earthworms), but only in the breeding season. Interestingly, we observed no increase in marine prey consumption after the landfill closures. In winter there was a decrease in terrestrial prey consumption, whereas the consumption of marine and, despite their reduce availability, landfill resources increased. Thus, and unlike when all the landfills were open, we detected significant trophic differences between breeding and non-breeding seasons. Additionally, we found significant trophic differences among colonies that were quite close to each other, but not between breeding adults and chicks. In conclusion, landfill closure or the use of deterrence systems had a direct impact on the trophic ecology of Yellow-legged Gulls; loss of refuse was mainly compensated for by prey of terrestrial origin when breeding, but not in winter. Thus, this species may experience foraging constraints in winter with potential effects on other life-history aspects including their dispersal, breeding and survival that needs further evaluation.     

Year‐round spatial movements and trophic ecology of Trindade Petrels (Pterodroma arminjoniana)

Trindade Petrels (Pterodroma arminjoniana) are vulnerable gadfly petrels that breed on the remote Trindade Island, located ~1100 km off the Brazilian coast. Little is known about their spatial ecology, and their trophic ecology has only been described for the breeding season. We tagged four Trindade Petrels with global location sensing loggers (GLS) from October 2013 to November 2014 and sampled the blood and feathers (innermost primary and the eighth secondary) of 14 individuals to evaluate their year‐round spatial and isotopic ecology. We examined individual distributions, habitat use and suitability, activity, and isotopic values during the breeding, migration, and non‐breeding periods. Trindade Petrels used areas in the southwest Atlantic Ocean (between 10°N and 50°S in latitude) during the breeding season. They migrated through pelagic waters of the tropical Atlantic to the northwest Atlantic, where they spent the non‐breeding season. Trindade Petrels used mostly tropical to subtropical waters in areas of intermediate to high wind speeds and low marine productivity. Individuals spent more time foraging at night than during the day. During the breeding season, birds in northerly areas had higher carbon‐13 values, and birds that used more pelagic areas foraged on prey at a higher trophic level (higher nitrogen‐15 values) than those in more southern and coastal areas. Isotopic values during the breeding, migration, and non‐breeding periods differed, possibly due to differences among individuals in their at‐sea distribution throughout the year. We confirmed the non‐breeding distribution of Trindade Petrels, which was previously known only from vessel sightings and stranded birds. Our results also suggest a strong temporal segregation in the at‐sea distribution and trophic ecology between two groups of individuals, which might indicate the existence of two separate breeding populations.     

A stable isotopic investigation into the causes of decline in a sub-Antarctic predator, the rockhopper penguin Eudyptes chrysocome

The rockhopper penguin (Eudyptes chrysocome) is a conspicuous apex marine predator that has experienced marked population declines throughout most of its circumpolar breeding distribution. The cause(s) for the declines remain elusive, but the relatively large spatio‐temporal scale over which population decreases have occurred implies that ecosystem‐scale, at‐sea factors are likely to be involved. We employ stable isotope analyses of carbon (¹³C/¹²C, expressed as δ¹³C) and nitrogen (¹⁵N/¹⁴N, δ¹⁵N) in time‐series of rockhopper penguin feather samples, dating back to 1861, in order to reconstruct the species' ecological history. Specifically, we examine whether rockhopper penguin population decline has been associated with a shift towards lower primary productivity in the ecosystem in which they feed, or with a shift to a diet of lower trophic status and lower quality, and we use long‐term temperature records to evaluate whether shifts in isotope ratios are associated with annual variations in sea surface temperature. Having controlled temporally for the Suess Effect and for increases in CO₂ concentrations in seawater, we found that overall, δ¹³C signatures decreased significantly over time in rockhopper penguins from seven breeding sites, supporting the hypothesis that decreases in primary productivity, and hence, carrying capacity, for which δ¹³C signature is a proxy, have been associated with the decline of penguin populations. There was some evidence of a long‐term decline in δ¹⁵N at some sites, and strong evidence that δ¹⁵N signatures were negatively related to sea surface temperatures across sites, indicative of a shift in diet to prey of lower trophic status over time and in warm years. However, a site‐by‐site analysis revealed divergent isotopic trends among sites: five of seven sites exhibited significant temporal or temperature‐related trends in isotope signatures. This study highlights the utility of stable isotope analyses when applied over relatively long timescales to apex predators.     

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.     

The Long and the Short of It: No Dietary Specialisation between Male and Female Western Sandpipers Despite Strong Bill Size Dimorphism

Many bird species show spatial or habitat segregation of the sexes during the non-breeding season. One potential ecological explanation is that differences in bill morphology favour foraging niche specialisation and segregation. Western sandpipers Calidris mauri have pronounced bill size dimorphism, with female bills averaging 15% longer than those of males. The sexes differ in foraging behaviour and exhibit partial latitudinal segregation during the non-breeding season, with males predominant in the north and females in the south. Niche specialisation at a local scale might account for this broad geographic pattern, and we investigated whether longer-billed females and shorter-billed males occupy different foraging niches at 16 sites across the non-breeding range. We used stable-nitrogen (δ¹⁵N) isotope analysis of whole blood to test for dietary specialisation according to bill length and sex. Stable-nitrogen isotope ratios increase with trophic level. We predicted that δ¹⁵N values would increase with bill length and would be higher for females, which use a greater proportion of foraging behaviour that targets higher-trophic level prey. We used stable-carbon (δ¹³C) isotope analysis to test for habitat segregation according to bill length and sex. Stable-carbon isotope ratios vary between marine- and freshwater-influenced habitats. We predicted that δ¹³C values would differ between males and females if the sexes segregate between habitat types. Using a model selection approach, we found little support for a relationship between δ¹⁵N and either bill length or sex. There was some indication, however, that more marine δ¹³C values occur with shorter bill lengths. Our findings provide little evidence that male and female western sandpipers exhibit dietary specialisation as a function of their bill size, but indicate that the sexes may segregate in different habitats according to bill length at some non-breeding sites. Potential ecological factors underlying habitat segregation between sexes include differences in preferred habitat type and predation risk.     

Population-Scale Foraging Segregation in an Apex Predator of the North Atlantic

In this work we investigated the between-colony spatial, behavioural and trophic segregation of two sub-populations of the elusive Macaronesian shearwaters Puffinus baroli breeding only ~340 km apart in Cima Islet (Porto Santo Island) and Selvagem Grande Island. Global location sensing (gls) loggers were used in combination with the trophic ecology of tracked individuals, inferred from the isotopic signatures of wing feathers. Results suggest that these two Macaronesian shearwater sub-populations do segregate during the non-breeding period in some ‘sub-population-specific’ regions, by responding to different oceanographic characteristics (habitat modelling). Within these disparate areas, both sub-populations behave differently (at-sea activity) and prey on disparate trophic niches (stable isotope analysis). One hypothesis would be that each sub-population have evolved and adapted to feed on particular and ‘sub-population-specific’ resources, and the segregation observed at the three different levels (spatial, behavioural and trophic) might be in fact a result of such adaptation, from the emergence of ‘cultural foraging patterns’. Finally, when comparing to the results of former studies reporting on the spatial, behavioural and trophic choices of Macaronesian shearwater populations breeding on Azores and Canary Islands, we realized the high ecological plasticity of this species inhabiting and foraging over the North-East Atlantic Ocean.     

Temporal variation in trophic relationships among three congeneric penguin species breeding in sympatry

Penguins are a monophyletic group in which many species are found breeding sympatrically, raising questions regarding how these species coexist successfully. Here, the isotopic niche of three sympatric pygoscelid penguin species was investigated at Powell Island, South Orkney Islands, during two breeding seasons (austral summers 2013–2014 and 2015–2016). Measurements of carbon (δ¹³C) and nitrogen (δ¹⁵N) stable isotope ratios were obtained from blood (adults) or feather (chicks) samples collected from Adélie Pygoscelis adeliae, chinstrap P. antarctica, and gentoo P. papua penguins. Isotopic niche regions (a proxy for the realized trophic niches) were computed to provide estimates of the trophic niche width of the studied species during the breeding season. The isotopic niche regions of adults of all three species were similar, but gentoo chicks had noticeably wider isotopic niches than the chicks of the other two species. Moderate to strong overlap in isotopic niche among species was found during each breeding season and for both age groups, suggesting that the potential for competition for shared food sources was similar during the two study years, although the actual level of competition could not be determined owing to the lack of data on resource abundance. Clear interannual shifts in isotopic niche were seen in all three species, though of lower amplitude for adult chinstrap penguins. These shifts were due to variation in carbon, but not nitrogen, isotopic ratios, which could indicate either a change in isotopic signature of their prey or a switch to an alternative food web. The main conclusions of this study are that (1) there is a partial overlap in the isotopic niches of these three congeneric species and that (2) they responded similarly to changes that likely occurred at the base of their food chain between the 2 years of the study.     

Stable isotope analysis as a tool to monitor dietary trends in little penguins Eudyptula minor

Long‐term dietary monitoring of seabirds can be used to relate population fluctuations to at‐sea events. Stomach flushing is a conventional dietary monitoring technique, but has a number of disadvantages. Stable isotope analysis (SIA) is a less invasive method that provides unbiased dietary information over a longer period. We evaluated stable isotope analysis as a potential tool for monitoring long‐term little penguin Eudyptula minor diet. We determined diet composition during the chick feeding stage using stomach flushing and SIA at three separate colonies, using spatial variation in diet as a surrogate for potential temporal variation. Bayesian isotopic mixing models were generated for blood and feathers to evaluate their ability to discriminate broad‐scale (fish, squid, crustaceans) and fine‐scale (individual prey species) diet composition. Differences in stable carbon and nitrogen isotope ratios were found between colonies: broad‐scale isotopic mixing models predicted different proportional contributions of broad taxa (fish, cephalopod, crustacean) to diet than was indicated by stomach samples, reflecting the bias incurred by one‐off stomach contents analysis. Fine‐scale isotopic mixing models predicted proportional contributions of prey items with less certainty. Blood isotopic mixing models had narrower confidence intervals than models for feathers, but trends in δ¹⁵N for feathers mirrored those for blood. Our results suggest that relying on stomach contents analysis to detect shifts in prey consumption in little penguins could be very misleading, resulting in a less‐than‐complete idea of total prey consumption. SIA of little penguin tissues could be used to monitor dietary shifts across dissimilar taxa that may affect population numbers, but would fail to detect shifts between fish species.     

Winter distribution and behaviour of gentoo penguins<Emphasis Type="Italic"> Pygoscelis papua</Emphasis> at South Georgia

Knowledge of the spatial and temporal dynamics of foraging penguins is important to our understanding of the Southern Ocean marine ecosystem. We use satellite tracking to provide the first data on the distribution and behaviour of gentoo penguins (Pygoscelis papua) during the winter at South Georgia. Five penguins tracked from Bird Island remained close inshore, and although they did not return to the initial tagging site, they did appear to return to land each evening. They made diurnal trips to sea of similar distance from land as those during the breeding season, even though the constraints of chick rearing were absent. Despite potential greater flexibility in their responses to variations in prey availability in winter, the penguins still returned to land each night. This may reflect benefits from conserving energy by resting on land, possibly facilitating information exchange and avoiding predation. The distribution and behaviour of gentoo penguins during the winter enables efficient exploitation of a dynamic, patchy prey resource and may ultimately determine the timing of return to the colony, and onset of breeding in the following season.     

Foraging behaviour of Magellanic Penguins during the early chick‐rearing period at Isla de los Estados,Argentina

Studies of the at‐sea distribution and trophic ecology of penguins are essential to understand their role in the broader marine food web. Magellanic Penguins Spheniscus magellanicus have a wide distribution and their foraging behaviour varies across breeding sites and between sexes, among others. In this study, we characterized the at‐sea areas, the diving strategies and the relative trophic level of Magellanic Penguins breeding at Isla de los Estados, Argentina, during the early chick‐rearing period. In addition, we quantified the interannual, sexual and individual variability in those parameters during three breeding seasons (2011–2013) using devices recording position and dive depth, and obtained blood samples for stable isotope analysis. During the early chick‐rearing period, Magellanic Penguins showed small differences between the sexes in their foraging behaviour and large overlap in the at‐sea areas used, suggesting no intraspecific variation between sexes. Although there was interannual variability in the foraging behaviour and the trophic level of the penguins, most of the studied nests managed successfully to raise both chicks during the first stage of the breeding cycle (guard stage). The foraging ecology of Magellanic Penguins from this colony was comparable with results of past studies at other breeding colonies. This study contributes to the identification of important at‐sea areas for Magellanic Penguins at the southern edge of their distribution and also to the identification of possible threats in the study area such as interaction with fisheries.     

Stable isotopes document the winter foraging ecology of king penguins and highlight connectivity between subantarctic and Antarctic ecosystems

The poorly known winter foraging ecology of the king penguin, a major Southern Ocean consumer, was investigated at the subantarctic Crozet Islands where the largest global population breeds. Blood δ¹³C and δ¹⁵N values were used as proxies of the birds’ foraging habitat and diet, respectively, and circulating prolactin levels helped in determining the birds’ reproductive status. Plasma prolactin concentrations showed that king penguin adults of unknown breeding status (n = 52) that were present at the colony in winter were in fact breeders and failed breeders, but were not non ‐breeders. Circulating prolactin was neither related to δ¹³C nor δ¹⁵N values, thus suggesting that both breeders and failed breeders used the same foraging habitats and fed on the same prey. Plasma and blood cell isotopic values depicted four new relevant biological features on the feeding strategies of king penguins during the critical winter period: (1) 42% of the birds foraged in the distant Antarctic Zone, but 58% fed primarily in subantarctic waters (δ¹³C), (2) they preyed upon myctophids in both zones (δ¹⁵N), (3) individuals were consistent in their foraging strategies over the winter months (δ¹³C and δ¹⁵N), and (4) a higher proportion of females (77%–80%) than males (27%–31%) favored feeding in distant Antarctic waters (δ¹³C). This study highlights trophic connectivity between subantarctic and Antarctic ecosystems and hence the key role of energy export from Antarctic waters to sustain breeding populations of subantarctic predators, including during the Austral winter.     

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.     

Using stable-isotope analysis of feathers to distinguish moulting and breeding origins of seabirds

To determine whether stable isotope measurements of bird feathers can be used to identify moulting (interbreeding) foraging areas of adult seabirds, we examined the stable-carbon (δ¹³C) and nitrogen (δ¹⁵N) isotopic composition of feathers of chicks and adults of black-browed albatrosses (Diomedea melanophrys) from Kerguelen Islands, southern Indian Ocean. Albatross chicks are fed primarily fish (75% by mass), the diet being dominated by various species of the family Nototheniidae and Channichthyidae which commonly occur in the shelf waters in the vicinity of the colony. δ¹³C and δ¹⁵N values in chick feathers, which are grown in summer in the breeding area, were lower than values in adult feathers, which are grown in winter (δ¹³C: –19.6‰ versus –17.6‰ and δ¹⁵N: 12.4‰ versus 15.7‰, respectively). No differences in δ¹³C and δ¹⁵N values were found in adult wing feathers moulted in 1993 and 1994 and in adult feathers formed at the beginning, middle and end of the 1994 moulting period. These data are consistent with adults moulting in the same area and feeding at the same trophic level from one year to the next and with no major changes in foraging ecology within a given moulting season; they suggest that foraging grounds were different in summer and winter and that these differed in their stable-isotope signature. Changes in both feather δ¹³C and δ¹⁵N values indicated feeding south of the Subtropical Front (STF) during chick rearing, which is in agreement with the known foraging ecology at this time and feeding north of the STF during moult. This, together with band recoveries from adult birds, indicates that black-browed albatrosses from Kerguelen Islands wintered in subtropical waters off southern Australia. The stable-isotope markers in feathers, therefore, have the potential for locating moulting areas of migratory seabird species moving between isotopically distinct regions and for investigating seabirds’ foraging ecology during the poorly known interbreeding period. Such information is needed for studies of year-round ecology of seabirds as well as for their conservation and the long-term monitoring of the pelagic environment. Received: 28 June 1999 / Accepted: 14 September 1999     

Variability in stable isotopes of snowy owl feathers and contribution of marine resources to their winter diet

The snowy owl is an elusive arctic predator known for its nomadic behaviour. Satellite tracking has revealed that some adult snowy owls could make an extensive use of the marine environment during the non‐breeding season. However, the relative contribution of marine resources to their diet is unknown. Stable isotope analyses can be useful to document the diet of mobile animals during periods of the year when individuals are less accessible. This study aimed to assess variation in isotopic values (δ¹³C and δ¹⁵N) of various feather types, and the usefulness of feathers to determine the contribution of the marine environment to the winter diet of snowy owls captured in summer. We sampled feathers coming from 6 body regions of 18 breeding females at two sites in the eastern Canadian Arctic in 2013 and 2014. Prior to analyses, diet‐tissue discrimination factors of snowy owl feathers were established in captivity. Variability in isotopic values among feather types was relatively low and pairwise correlations in isotopic values between feathers on the same individual were variable and often low, which suggests differences in the diet at the time when various feathers were synthesized. Diet reconstruction models detected a contribution of marine sources to snowy owl feathers ranging from 4 to 19% among feather types. However, the marine contribution was highly variable when single feathers were examined within individuals, ranging from 3 to 71%. This indicated that no single feather type could be used alone to reliably infer the contribution of marine resources to the winter diet of owls, possibly due to a high variability in the timing and sequence of molt. For asynchronous molters like snowy owls, we recommend sampling multiple feathers from various body regions, excluding wing feathers, to investigate winter diet or habitat use.     

Daily activity and minimum food requirements during winter for gentoo penguins (Pygoscelis papua) in the South Shetland Islands, Antarctica

Estimates of daily activity and consequent demand for food during winter are scarce for many polar seabirds, yet essential for assessing constraints on foraging effort, demand for food, and potential competition with local fisheries. We affixed archival temperature tags to gentoo penguins (Pygoscelis papua) from two colonies in the South Shetland Islands to measure the frequency, timing, and duration of foraging trips and to estimate minimum food requirements during winter. Foraging trip frequencies ranged from 0.85 to 1.0 trips day⁻¹ and were positively correlated with day length. Early winter foraging trips more closely matched day length than late winter foraging trips. The data suggest that individuals maximize foraging time during the early winter period, likely to recover body mass following the breeding season and molt. The more attenuated response of foraging trip durations to increasing day length in late winter may be related to differences in local resource availability or individual behaviors prior to the upcoming breeding season. Minimum food requirements also exhibited a seasonal cycle with a mid-winter minimum. On average, minimum food requirements were estimated at 0.70 ± 0.12 kg day⁻¹. Extrapolated to the regional population of gentoo penguins, winter food requirements by gentoo penguins were equivalent to roughly 33% of annual krill catches by commercial fisheries in the South Shetland Island region over the past decade. Current expansion of the gentoo population and the krill fishery in the southern Scotia Sea warrants continued monitoring of gentoo penguins during winter.