Michael J. Polito

Integrating Stomach Content and Stable Isotope Analyses to Quantify the Diets of Pygoscelid Penguins

Stomach content analysis (SCA) and more recently stable isotope analysis (SIA) integrated with isotopic mixing models have become common methods for dietary studies and provide insight into the foraging ecology of seabirds. However, both methods have drawbacks and biases that may result in difficulties in quantifying inter-annual and species-specific differences in diets. We used these two methods to simultaneously quantify the chick-rearing diet of Chinstrap (Pygoscelis antarctica) and Gentoo (P. papua) penguins and highlight methods of integrating SCA data to increase accuracy of diet composition estimates using SIA. SCA biomass estimates were highly variable and underestimated the importance of soft-bodied prey such as fish. Two-source, isotopic mixing model predictions were less variable and identified inter-annual and species-specific differences in the relative amounts of fish and krill in penguin diets not readily apparent using SCA. In contrast, multi-source isotopic mixing models had difficulty estimating the dietary contribution of fish species occupying similar trophic levels without refinement using SCA-derived otolith data. Overall, our ability to track inter-annual and species-specific differences in penguin diets using SIA was enhanced by integrating SCA data to isotopic mixing modes in three ways: 1) selecting appropriate prey sources, 2) weighting combinations of isotopically similar prey in two-source mixing models and 3) refining predicted contributions of isotopically similar prey in multi-source models.

A 1000-year record of Adelie penguin diets in the Southern Ross Sea

Non-krill prey remains were recovered from ornithogenic sediments at three active Adélie penguin colonies on Ross Island, to assess long-term dietary trends in this species. Radiocarbon dates place the age of these deposits from a maximum of 947 years ago to the present. We identified 12 taxa of fish and two of squid with the Antarctic silverfish (Pleuragramma antarcticum) as the most abundant prey species represented at all sites. In addition, silverfish have decreased in importance in Adélie penguin diet over the past 600 years, perhaps in response to climate change since the onset of the Little Ice Age, though it remains much more abundant in current penguin diet in the Ross Sea than in the Antarctic Peninsula. Other prey taxa reflect the diversity of prey selection by Adélie penguins in Antarctica.

Carbon and nitrogen isotope fractionation of amino acids in an avian marine predator, the gentoo penguin (Pygoscelis papua)

Compound-specific stable isotope analysis (CSIA) of amino acids (AA) has rapidly become a powerful tool in studies of food web architecture, resource use, and biogeochemical cycling. However, applications to avian ecology have been limited because no controlled studies have examined the patterns in AA isotope fractionation in birds. We conducted a controlled CSIA feeding experiment on an avian species, the gentoo penguin (Pygoscelis papua), to examine patterns in individual AA carbon and nitrogen stable isotope fractionation between diet (D) and consumer (C) (Δ13CC-D and Δ15NC-D, respectively). We found that essential AA δ13C values and source AA δ15N values in feathers showed minimal trophic fractionation between diet and consumer, providing independent but complimentary archival proxies for primary producers and nitrogen sources respectively, at the base of food webs supporting penguins. Variations in nonessential AA Δ13CC-D values reflected differences in macromolecule sources used for biosynthesis (e.g., protein vs. lipids) and provided a metric to assess resource utilization. The avian-specific nitrogen trophic discrimination factor (TDFGlu-Phe = 3.5 ± 0.4‰) that we calculated from the difference in trophic fractionation (Δ15NC-D) of glutamic acid and phenylalanine was significantly lower than the conventional literature value of 7.6‰. Trophic positions of five species of wild penguins calculated using a multi-TDFGlu-Phe equation with the avian-specific TDFGlu-Phe value from our experiment provided estimates that were more ecologically realistic than estimates using a single TDFGlu-Phe of 7.6‰ from the previous literature. Our results provide a quantitative, mechanistic framework for the use of CSIA in nonlethal, archival feathers to study the movement and foraging ecology of avian consumers.

A reversal of fortunes: climate change ‘winners’ and ‘losers’ in Antarctic Peninsula penguins

Climate change is a major threat to global biodiversity. Antarctic ecosystems are no exception. Investigating past species responses to climatic events can distinguish natural from anthropogenic impacts. Climate change produces ‘winners’, species that benefit from these events and ‘losers’, species that decline or become extinct. Using molecular techniques, we assess the demographic history and population structure of Pygoscelis penguins in the Scotia Arc related to climate warming after the Last Glacial Maximum (LGM). All three pygoscelid penguins responded positively to post-LGM warming by expanding from glacial refugia, with those breeding at higher latitudes expanding most. Northern (Pygoscelis papua papua) and Southern (Pygoscelis papua ellsworthii) gentoo sub-species likely diverged during the LGM. Comparing historical responses with the literature on current trends, we see Southern gentoo penguins are responding to current warming as they did during post-LGM warming, expanding their range southwards. Conversely, Adélie and chinstrap penguins are experiencing a ‘reversal of fortunes’ as they are now declining in the Antarctic Peninsula, the opposite of their response to post-LGM warming. This suggests current climate warming has decoupled historic population responses in the Antarctic Peninsula, favoring generalist gentoo penguins as climate change ‘winners’, while Adélie and chinstrap penguins have become climate change ‘losers’.

Integrating oxidative ecology into conservation physiology.

Given that oxidative balance acts on fitness components, its measurement may be valuable to conservationists to assess population health. We show that antioxidant defences reflect population trends in penguin colonies. These preliminary results suggest that oxidative balance could be used to assess the health of animal populations in their habitat.

Spatial and isotopic niche partitioning during winter in chinstrap and Adélie penguins from the South Shetland Islands

Closely related species with similar ecological requirements should exhibit segregation along spatial, temporal, or trophic niche axes to limit the degree of competitive overlap. For migratory marine organisms like seabirds, assessing such overlap during the non-breeding period is difficult because of long-distance dispersal to potentially diffuse foraging habitats. Miniaturization of geolocation devices and advances in stable isotope analysis (SIA), however, provide a robust toolset to quantitatively track the movements and foraging niches of wide ranging marine animals throughout much of their annual cycle. We used light-based geolocation tags and analyzed stable carbon and nitrogen isotopes from tail feathers to simultaneously characterize winter movements, habitat utilization, and overlap of spatial and isotopic niches of migratory chinstrap (Pygoscelis antarctica) and Adelie (P. adeliae) penguins during the austral winter of 2012. Chinstrap penguins exhibited a higher diversity of movements and occup...

Penguin eggshell membranes reflect homogeneity of mercury in the marine food web surrounding the Antarctic Peninsula.

Remote regions such as the Antarctic have become increasingly important for investigations into far-reaching anthropogenic impacts on the environment, most recently in regard to the global mercury cycle. Spatial patterns of mercury availability in four regions of the Antarctic Peninsula were investigated using three species of sympatrically breeding Pygoscelis penguins as biomonitors. Eggshells with intact membranes from Adélie, Gentoo, and Chinstrap penguins were collected at 24 breeding colonies in the South Orkney Islands, South Shetland Islands, eastern Antarctic Peninsula, and western Antarctic Peninsula during the 2006/2007 austral summer. In addition, we compared eggshell membrane mercury concentrations with eggshell stable isotope values (δ(15)N and δ(13)C) to determine if species-specific trophic or foraging habitat preferences influenced female mercury exposure prior to breeding. With few exceptions, mercury concentrations were found to be fairly homogeneous throughout the Antarctic Peninsula suggesting little spatial variation in the risk of exposure to dietary mercury in this food web. Mercury concentrations in Gentoo and Adélie penguins were similar while Chinstrap penguins tended to have higher eggshell membrane mercury concentrations than their congeners. However, inter and intra-specific differences in eggshell membrane mercury concentration were not related to eggshell δ(15)N or δ(13)C values, a likely result of all three species foraging at similar trophic positions. The lack of regional-scale differences in mercury availability in this marine ecosystem may be a reflection of generally uniform atmospheric deposition and upwelling of regionally homogeneous deep water rather than from geographically distinct point sources.

Trophic calculations reveal the mechanism of population-level variation in mercury concentrations between marine ecosystems: case studies of two polar seabirds.

The incorporation of quantitative trophic level analysis in ecotoxicological studies provides explanatory power to identify the factors, trophic or environmental, driving population-level variation in mercury exposure at large geographic scales. In the Antarctic marine ecosystem, mercury concentrations and stable isotope values in Adélie penguins (Pygoscelis adeliae) were compared between the Antarctic Peninsula and the Ross Sea. Correcting tissue δ(15)N values for baseline δ(15)N values revealed population-level differences in trophic position which contributes to differences in mercury. Data from Thick-billed murres (Uria lomvia) were synthesized from published values from Baffin Bay and Svalbard to demonstrate the utility of baseline δ(15)N values in identifying differences in environmental mercury exposure independent of diet. Here, we demonstrate the importance of calculating population-specific trophic level data to uncover the source of variation in mercury concentrations between geographically distinct populations of marine predators.

Multi-tissue analyses reveal limited inter-annual and seasonal variation in mercury exposure in an Antarctic penguin community

Inter-annual variation in tissue mercury concentrations in birds can result from annual changes in the bioavailability of mercury or shifts in dietary composition and/or trophic level. We investigated potential annual variability in mercury dynamics in the Antarctic marine food web using Pygoscelis penguins as biomonitors. Eggshell membrane, chick down, and adult feathers were collected from three species of sympatrically breeding Pygoscelis penguins during the austral summers of 2006/2007–2010/2011. To evaluate the hypothesis that mercury concentrations in penguins exhibit significant inter-annual variation and to determine the potential source of such variation (dietary or environmental), we compared tissue mercury concentrations with trophic levels as indicated by δ15N values from all species and tissues. Overall, no inter-annual variation in mercury was observed in adult feathers suggesting that mercury exposure, on an annual scale, was consistent for Pygoscelis penguins. However, when examining tissues that reflected more discrete time periods (chick down and eggshell membrane) relative to adult feathers, we found some evidence of inter-annual variation in mercury exposure during penguins’ pre-breeding and chick rearing periods. Evidence of inter-annual variation in penguin trophic level was also limited suggesting that foraging ecology and environmental factors related to the bioavailability of mercury may provide more explanatory power for mercury exposure compared to trophic level alone. Even so, the variable strength of relationships observed between trophic level and tissue mercury concentrations across and within Pygoscelis penguin species suggest that caution is required when selecting appropriate species and tissue combinations for environmental biomonitoring studies in Antarctica.

Unique pattern of molt leads to low intraindividual variation in feather mercury concentrations in penguins

The authors hypothesized that the catastrophic annual molt of penguins (Sphenisciformes) would lead to reduced intraindividual variation of mercury concentrations in body feathers. While mean mercury concentrations varied significantly among 8 penguin species, intraindividual variability did not differ among species and was 3 times lower than values observed in other seabirds. The findings of the present study suggest that a single body feather collected at random per individual can be adequate to estimate mercury exposure at the population level in penguins.