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If not carefully managed, harvesting of krill risks disturbing the ecological balance of many Antarctic and sub-Antarctic sites where krill-dependent predators feed. One of the least disturbed sites anywhere within the Southern Ocean and one where krill fishing has so far been virtually non-existent is the South Sandwich Islands volcanic archipelago. Some of the main krill predators breeding at the South Sandwich Islands are penguins, with five species breeding on the islands, the dominant species of which is the chinstrap penguin. In this paper we report on the results of ARGOS PTT deployments during the chinstrap penguin chick-rearing period, using the recorded foraging trips to develop habitat models. Foraging habitats used by chinstrap penguins during the chick-rearing period were best characterised by distance from the colony and sea surface temperature and, using these two covariates, we predicted foraging habitat use around all islands. We show that the provisions of the South Georgia and South Sandwich Islands Marine Protected Area ensure that chinstrap penguins, and other krill-dependent predators with similar foraging ranges, likely have robust protection during the summer. During the winter, when krill predators are likely to forage further offshore, seasonal sea ice provides a physical barrier to exclude the fishery, again ensuring the islands’ unique biodiversity receives strong protection. However, to the north of the marginal sea ice zone, competition between krill predators and the fishery could exist if the fishery were ever to explore new locations for resource extraction. We make a number of conclusions, including the need for winter tracking data to inform future management options.

Many remote islands present barriers to effective wildlife monitoring in terms of challenging terrain and frequency of visits. The sub-Antarctic islands of South Georgia and the South Sandwich Islands are home to globally significant populations of seabirds and marine mammals. South Georgia hosts the largest breeding populations of Antarctic fur seals, southern elephant seals and king penguins as well as significant populations of wandering, black-browed and grey-headed albatross. The island also holds important populations of macaroni and gentoo penguins. The South Sandwich Islands host the world’s largest colony of chinstrap penguins in addition to major populations of Adélie and macaroni penguins. A marine protected area was created around these islands in 2012 but monitoring populations of marine predators remains a challenge, particularly as these species breed over large areas in remote and often inaccessible locations. During the 2019/20 austral summer, we trialed the use of an unoccupied aerial vehicle (UAV; drone) to monitor populations of seals, penguins and albatross and here we report our initial findings, including considerations about the advantages and limitations of the methodology. Three extensive southern elephant seal breeding sites were surveyed with complete counts made around the peak pupping date, two of these sites were last surveyed 24 years ago. A total of nine islands, historically recorded as breeding sites for wandering albatross, were surveyed with 144 fledglings and 48 adults identified from the aerial imagery. The UAV was effective at surveying populations of penguins that nest on flat, open terrain, such as Adélie and chinstrap penguin colonies at the South Sandwich Islands, and an extensive king penguin colony on South Georgia, but proved ineffective for monitoring macaroni penguins nesting in tussock habitat on South Georgia as individuals were obscured or hidden by vegetation. Overall, we show that UAV surveys can allow regular an

© The Author(s), 2020. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Trathan, P. N., Wienecke, B., Barbraud, C., Jenouvrier, S., Kooyman, G., Le Bohec, C., Ainley, D. G., Ancel, A., Zitterbart, D. P., Chown, S. L., LaRue, M., Cristofari, R., Younger, J., Clucas, G., Bost, C., Brown, J. A., Gillett, H. J., & Fretwell, P. T. The emperor penguin - vulnerable to projected rates of warming and sea ice loss. Biological Conservation, 241, (2020): 108216, doi:10.1016/j.biocon.2019.108216., We argue the need to improve climate change forecasting for ecology, and importantly, how to relate long-term projections to conservation. As an example, we discuss the need for effective management of one species, the emperor penguin, Aptenodytes forsteri. This species is unique amongst birds in that its breeding habit is critically dependent upon seasonal fast ice. Here, we review its vulnerability to ongoing and projected climate change, given that sea ice is susceptible to changes in winds and temperatures. We consider published projections of future emperor penguin population status in response to changing environments. Furthermore, we evaluate the current IUCN Red List status for the species, and recommend that its status be changed to Vulnerable, based on different modelling projections of population decrease of ≥50% over the current century, and the specific traits of the species. We conclude that current conservation measures are inadequate to protect the species under future projected scenarios. Only a reduction in anthropogenic greenhouse gas emissions will reduce threats to the emperor penguin from altered wind regimes, rising temperatures and melting sea ice; until such time, other conservation actions are necessary, including increased spatial protection at breeding sites and foraging locations. The designation of large-scale marine spatial protection across its range would benefit the species, particularly in areas that have a high probability of becoming future climate change refugia. We also recommend that the emperor penguin is listed by the Antarctic Treaty as an Antarctic Specially Protected Species, with development of a species Action Plan., We thank Thomas J. Bracegirdle, Tony Phillips and Kevin Hughes for helpful comments on earlier drafts of this manuscript. PNT acknowledges the support of WWF-UK under GB095701 and SJ the support of NSF OPP1744794 and 1643901.

Over evolutionary time, pathogen challenge shapes the immune phenotype of the host to better respond to an incipient threat. The extent and direction of this selection pressure depends on the local pathogen composition, which is in turn determined by biotic and abiotic features of the environment. However, little is known about adaptation to local pathogen threats in wild animals. The Gentoo penguin (Pygoscelis papua) is a species complex that lends itself to the study of immune adaptation because of its circumpolar distribution over a large latitudinal range, with little or no admixture between different clades. In this study, we examine the diversity in a key family of innate immune genes - the Toll-like receptors (TLRs) - across the range of the Gentoo. The three TLRs that we investigated present varying levels of diversity, with TLR4 and TLR5 greatly exceeding the diversity of TLR7. We present evidence of positive selection in TLR4 and TLR5, which points to pathogen-driven adaptation to the local pathogen milieu. Finally, we demonstrate that two positively selected co-segregating sites in TLR5 are sufficient to alter the responsiveness of the receptor to its bacterial ligand, flagellin. Taken together, these results suggest that Gentoo penguins have experienced distinct pathogen-driven selection pressures in different environments, which may be important given the role of the Gentoo as a sentinel species in some of the world’s most rapidly changing environments.

We argue the need to improve climate change forecasting for ecology, and importantly, how to relate long-term projections to conservation. As an example, we discuss the need for effective management of one species, the emperor penguin, Aptenodyptes forsteri. This species is unique amongst birds in that its breeding habit is critically dependent upon seasonal fast ice. Here, we review its vulnerability to ongoing and projected climate change, given that sea ice is susceptible to changes in winds and temperatures. We consider published projections of future emperor penguin population status in response to changing environments. Furthermore, we evaluate the current IUCN Red List status for the species, and recommend that its status be changed to Vulnerable, based on different modelling projections of population decrease of ≥50% over the current century, and the specific traits of the species. We conclude that current conservation measures are inadequate to protect the species under future projected scenarios. Only a reduction in anthropogenic greenhouse gas emissions will reduce threats to the emperor penguin from altered wind regimes, rising temperatures and melting sea ice; until such time, other conservation actions are necessary, including increased spatial protection at breeding sites and foraging locations. The designation of large-scale marine spatial protection across its range would benefit the species, particularly in areas that have a high probability of becoming future climate change refugia. We also recommend that the emperor penguin is listed by the Antarctic Treaty as an Antarctic Specially Protected Species, with development of a species Action Plan.

Over evolutionary time, pathogen challenge shapes the immune phenotype of the host to better respond to an incipient threat. The extent and direction of this selection pressure depend on the local pathogen composition, which is in turn determined by biotic and abiotic features of the environment. However, little is known about adaptation to local pathogen threats in wild animals. The Gentoo penguin (Pygoscelis papua) is a species complex that lends itself to the study of immune adaptation because of its circumpolar distribution over a large latitudinal range, with little or no admixture between different clades. In this study, we examine the diversity in a key family of innate immune genes-the Toll-like receptors (TLRs)-across the range of the Gentoo penguin. The three TLRs that we investigated present varying levels of diversity, with TLR4 and TLR5 greatly exceeding the diversity of TLR7. We present evidence of positive selection in TLR4 and TLR5, which points to pathogen-driven adaptation to the local pathogen milieu. Finally, we demonstrate that two positively selected cosegregating sites in TLR5 are sufficient to alter the responsiveness of the receptor to its bacterial ligand, flagellin. Taken together, these results suggest that Gentoo penguins have experienced distinct pathogen-driven selection pressures in different environments, which may be important given the role of the Gentoo penguin as a sentinel species in some of the world's most rapidly changing environments.

© The Author(s), 2018. This article is distributed under the terms of the Creative Commons Attribution License. The definitive version was published in Scientific Reports 8 (2018): 3926, doi:10.1038/s41598-018-22313-w., Despite concerted international effort to track and interpret shifts in the abundance and distribution of Adélie penguins, large populations continue to be identified. Here we report on a major hotspot of Adélie penguin abundance identified in the Danger Islands off the northern tip of the Antarctic Peninsula (AP). We present the first complete census of Pygoscelis spp. penguins in the Danger Islands, estimated from a multi-modal survey consisting of direct ground counts and computer-automated counts of unmanned aerial vehicle (UAV) imagery. Our survey reveals that the Danger Islands host 751,527 pairs of Adélie penguins, more than the rest of AP region combined, and include the third and fourth largest Adélie penguin colonies in the world. Our results validate the use of Landsat medium-resolution satellite imagery for the detection of new or unknown penguin colonies and highlight the utility of combining satellite imagery with ground and UAV surveys. The Danger Islands appear to have avoided recent declines documented on the Western AP and, because they are large and likely to remain an important hotspot for avian abundance under projected climate change, deserve special consideration in the negotiation and design of Marine Protected Areas in the region., We gratefully acknowledge the financial support of the Dalio Foundation, Inc. through the Dalio Explore Fund, which provided all the financing for the Danger Island Expedition. We would like to thank additional support for analysis from the National Science Foundation (NSF PLR&GSS 1255058 - H.J.L. and P.M.; NSF PLR 1443585 – M.J.P.) and the National Aeronautical and Space Administration (NNX14AC32G; H.J.L. and M.S.). Geospatial support for the analysis of high resolution satellite imagery provided by the Polar Geospatial Center under NSF PLR awards 1043681 & 1559691.

The mechanisms that determine patterns of species dispersal are important factors in the production and maintenance of biodiversity. Understanding these mechanisms helps to forecast the responses of species to environmental change. Here we used a comparative framework and genome‐wide data obtained through RAD‐seq to compare the patterns of connectivity among breeding colonies for five penguin species with shared ancestry, overlapping distributions, and differing ecological niches, allowing an examination of the intrinsic and extrinsic barriers governing dispersal patterns. Our findings show that at‐sea range and oceanography underlie patterns of dispersal in these penguins. The pelagic niche of emperor (Aptenodytes forsteri), king (A. patagonicus), Adélie (Pygoscelis adeliae) and chinstrap (P. antarctica) penguins facilitates gene flow over thousands of kilometres. In contrast, the coastal niche of gentoo penguins (P. papua) limits dispersal, resulting in population divergences. Oceanographic fronts also act as dispersal barriers to some extent. We recommend that forecasts of extinction risk incorporate dispersal and that management units are defined by at‐sea range and oceanography in species lacking genetic data.

Climate change, fisheries' pressure on penguin prey, and direct human disturbance of wildlife have all been implicated in causing large shifts in the abundance and distribution of penguins in the Southern Ocean. Without mark-recapture studies, understanding how colonies form and, by extension, how ranges shift is challenging. Genetic studies, particularly focused on newly established colonies, provide a snapshot of colonization and can reveal the extent to which shifts in abundance and occupancy result from changes in demographic rates (e.g., reproduction and survival) or migration among suitable patches of habitat. Here, we describe the population structure of a colonial seabird breeding across a large latitudinal range in the Southern Ocean. Using multilocus microsatellite genotype data from 510 Gentoo penguin (Pygoscelis papua) individuals from 14 colonies along the Scotia Arc and Antarctic Peninsula, together with mitochondrial DNA data, we find strong genetic differentiation between colonies north and south of the Polar Front, that coincides geographically with the taxonomic boundary separating the subspecies P. p. papua and P. p. ellsworthii. Using a discrete Bayesian phylogeographic approach, we show that southern Gentoos expanded from a possible glacial refuge in the center of their current range, colonizing regions to the north and south through rare, long-distance dispersal. Our findings show that this dispersal is important for new colony foundation and range expansion in a seabird species that ordinarily exhibits high levels of natal philopatry, though persistent oceanographic features serve as barriers to movement.

The relationship between population structure and demographic history is critical to understanding microevolution and for predicting the resilience of species to environmental change. Using mitochondrial DNA from extant colonies and radiocarbon-dated subfossils, we present the first microevolutionary analysis of emperor penguins (Aptenodytes forsteri) and show their population trends throughout the last glacial maximum (LGM, 19.5–16 kya) and during the subsequent period of warming and sea ice retreat. We found evidence for three mitochondrial clades within emperor penguins, suggesting that they were isolated within three glacial refugia during the LGM. One of these clades has remained largely isolated within the Ross Sea, while the two other clades have intermixed around the coast of Antarctica from Adélie Land to the Weddell Sea. The differentiation of the Ross Sea population has been preserved despite rapid population growth and opportunities for migration. Low effective population sizes during the LGM, followed by a rapid expansion around the beginning of the Holocene, suggest that an optimum set of sea ice conditions exist for emperor penguins, corresponding to available foraging area.