Delay in Adélie penguin nest occupation restricts parental investment in nest construction and reduces reproductive output.

Reproductive success is an important demographic parameter that can be driven by environmental and behavioural factors operating on various spatio-temporal scales. As seabirds breed on land and forage in the ocean, processes occurring in both environments can influence their reproductive success. At various locations around East Antarctica, Adélie penguins' (Pygoscelis adeliae) reproductive success has been negatively linked to extensive sea-ice. In contrast, our study site in the Windmill Islands has limited fast ice present during the breeding season, allowing us to examine drivers of reproductive success under vastly different marine environmental conditions. Here, we examined the reproductive success of 450 Adélie penguin nests over a 10-year period using images obtained from remotely operated cameras. We analysed nest survival in relation to marine and climatic factors, environmental conditions at the camera site and immediately around the nest, and behavioural attributes reflecting parental investment and phenological timing. Our key result was a strong positive association between nest structure and chick survival, particularly when ground moisture and snow cover around the nest were high. Earlier nesting birds were more likely to build bigger nests, although it is unclear whether this is due to more time available to build nests or whether early arrival and high-quality nests are complementary traits. This intrinsic activity is likely to become more important if future predictions of increased snowfall in this region manifest.

Circumpolar assessment of mercury contamination: the Adélie penguin as a bioindicator of Antarctic marine ecosystems.

Due to its persistence and potential ecological and health impacts, mercury (Hg) is a global pollutant of major concern that may reach high concentrations even in remote polar oceans. In contrast to the Arctic Ocean, studies documenting Hg contamination in the Southern Ocean are spatially restricted and large-scale monitoring is needed. Here, we present the first circumpolar assessment of Hg contamination in Antarctic marine ecosystems. Specifically, the Adélie penguin (Pygoscelis adeliae) was used as a bioindicator species, to examine regional variation across 24 colonies distributed across the entire Antarctic continent. Mercury was measured on body feathers collected from both adults (n = 485) and chicks (n = 48) between 2005 and 2021. Because penguins' diet represents the dominant source of Hg, feather δ13C and δ15N values were measured as proxies of feeding habitat and trophic position. As expected, chicks had lower Hg concentrations (mean ± SD: 0.22 ± 0.08 µg·g‒1) than adults (0.49 ± 0.23 µg·g‒1), likely because of their shorter bioaccumulation period. In adults, spatial variation in feather Hg concentrations was driven by both trophic ecology and colony location. The highest Hg concentrations were observed in the Ross Sea, possibly because of a higher consumption of fish in the diet compared to other sites (krill-dominated diet). Such large-scale assessments are critical to assess the effectiveness of the Minamata Convention on Mercury. Owing to their circumpolar distribution and their ecological role in Antarctic marine ecosystems, Adélie penguins could be valuable bioindicators for tracking spatial and temporal trends of Hg across Antarctic waters in the future.

Environment-triggered demographic changes cascade and compound to propel a dramatic decline of an Antarctic seabird metapopulation.

While seabirds are well-known for making a living under some of the harshest conditions on the planet, their capacity to buffer against unfavourable conditions can be stretched in response to ecosystem change. During population increases, overlap between conspecifics can limit population growth through competition for breeding or feeding resources. What is less well understood is the role that intrinsic processes play during periods of population decline or under a changing environment. We interrogate key demographic parameters and their biophysical drivers to understand the role of intrinsic and extrinsic drivers during a recent near halving of a large Adélie penguin (Pygoscelis adeliae) metapopulation. The loss of 154,000 breeding birds along the 100-km East Antarctic coastline centred around 63°E over the last decade diverges from a sustained increase over preceding decades and is contrary to recent models that predict a continued increase. The decline was initially triggered by changed environmental conditions: more extensive near-shore sea ice caused a reduction in breeding success. The evidence suggests this decline was exacerbated by feedback processes driving an inverse density-dependent decrease in fledgling survival in response to smaller cohort size. It appears that the old adage of safety in numbers may shape the fledgling penguins' chances of survival and, if compromised over multiple years, could exacerbate difficulties during population decline or if feedback processes arise. The likely interplay between demographic parameters meant that conditions were more unfavourable and negative effects more rapid than would be expected if demographic processes acted in isolation or independently. Failure to capture both intrinsic and extrinsic drivers in predictive population models may mean that the real impacts of climate change on species' populations are more severe than projections would lead us to believe. These results improve our understanding of population regulation during periods of rapid decline for long-lived marine species.

Phenology-based adjustments improve population estimates of Antarctic breeding seabirds: the case of Cape petrels in East Antarctica.

To monitor and conserve a species, it is crucial to understand the size and distribution of populations. For seabirds, population surveys are usually conducted at peak breeding attendance. One of the largest populations of Cape petrels in East Antarctica is at the Vestfold Islands, where environmental and logistical constraints often prevent access to breeding sites at the optimal time for population surveys. In this study, we aim to quantify the contemporary and historical breeding population size of these Cape petrels by adjusting nest counts for variation in breeding phenology using photographs from remote cameras. We also compare spatial distribution between 1970s and 2017/2018. Our results show ground counts occurred outside peak breeding attendance, and adjusting for phenology changed the contemporary and historical population estimates. The Cape petrels showed local intra-island or adjacent-island changes in their distribution between the 1970s and 2017/2018 with no evidence of expanding or restricting their distribution or a significant change in their breeding population size. The results emphasize the importance of accounting for phenology in population counts, where populations are inaccessible at an optimal survey time. We discuss the applications of our research methodology for populations breeding in remote areas and as a baseline for assessing population change.

South polar skua (Catharacta maccormicki) as biovectors for long-range transport of persistent organic pollutants to Antarctica.

Migratory bird species may serve as vectors of contaminants to Antarctica through the local deposition of guano, egg abandonment, or mortality. To further investigate this chemical input pathway, we examined the contaminant burdens and profiles of the migratory South polar skua (Catharacta maccormicki) and compared them to the endemic Adélie penguin (Pygoscelis adeliae). A range of persistent organic pollutants were targeted in muscle and guano to facilitate differentiation of likely exposure pathways. A total of 56 of 65 targeted analytes were detected in both species, but there were clear profile and magnitude differences between the species. The South polar skua and Adélie penguin muscle tissue burdens were dominated by p,p'-dichlorodiphenyldichloroethylene (mean 5600 ng g-1 lw and 330 ng g-1 lw respectively) and hexachlorobenzene (mean 2500 ng g-1 lw and 570 ng g-1 lw respectively), a chemical profile characteristic of the Antarctic and Southern Ocean region. Species profile differences, indicative of exposure at different latitudes, were observed for polychlorinated biphenyls (PCBs), with lower chlorinated congeners and deca-chlorinated PCB-209 detected in South polar Skua, but not in Adélie penguins. Notably, the more recently used perfluoroalkyl substances and the brominated flame retardants, hexabromocyclododecane and tetrabromobisphenol A, were detected in both species. This finding suggests local exposure, given the predicted slow and limited long-range environmental transport capacity of these compounds to the eastern Antarctic sector.

Emerging evidence of resource limitation in an Antarctic seabird metapopulation after 6 decades of sustained population growth.

The influence of resource limitation on spatio-temporal population dynamics is a fundamental theme in ecology and the concepts of carrying capacity, density dependence and population synchrony are central to this theme. The life history characteristics of seabirds, which include use of disjunct patches of breeding habitat, high coloniality during breeding, strong philopatry, and central-place foraging, make this group well suited to studying this paradigm. Here, we investigate whether density-dependent processes are starting to limit population growth in the Adélie penguin metapopulation breeding in the Windmill Islands, East Antarctica, after 6 decades of growth. Our finding that the regional growth rate has slowed in recent decades, and that growth is slowing differentially across local populations as availability of breeding habitat and possibly food resources decrease, supports the notion of density-dependent regulation. Our observation of the first new colonisation of a breeding patch in a half-century of population growth by this highly philopatric species is further evidence for this. Given these emerging patterns of spatio-temporal population dynamics, this metapopulation may be at a point where the rate of change in density-dependent processes and rare events such as colonisations accelerates into the future, potentially providing new insights into spatio-temporal metapopulation dynamics of a long-lived species over a short time-frame. Continued long-term study of populations experiencing these circumstances provides an opportunity to expedite advances in understanding metapopulation processes. Our study highlights the importance of spatial heterogeneity and the mosaic of abiotic and biotic features of landscapes and seascapes in shaping species' metapopulation dynamics.

Density dependence forces divergent population growth rates and alters occupancy patterns of a central place foraging Antarctic seabird.

Density-dependent regulation is an important process in spatio-temporal population dynamics because it can alter the effects of synchronizing processes operating over large spatial scales. Most frequently, populations are regulated by density dependence when higher density leads to reduced individual fitness and population growth, but inverse density dependence can also occur when small populations are subject to higher extinction risks. We investigate whether density-dependent regulation influences population growth for the Antarctic breeding Adélie penguin Pygoscelis adeliae. Understanding the prevalence and nature of density dependence for this species is important because it is considered a sentinel species reflecting the impacts of fisheries and environmental change over large spatial scales in the Southern Ocean, but the presence of density dependence could introduce uncertainty in this role. Using data on population growth and indices of resource availability for seven regional Adélie penguin populations located along the East Antarctic coastline, we find compelling evidence that population growth is constrained at some locations by the amount of breeding habitat available to individuals. Locations with low breeding habitat availability had reduced population growth rates, higher overall occupancy rates, and higher occupancy of steeper slopes that are sparsely occupied or avoided at other locations. Our results are consistent with evolutionary models of avian breeding habitat selection where individuals search for high-quality nest sites to maximize fitness returns and subsequently occupy poorer habitat as population density increases. Alternate explanations invoking competition for food were not supported by the available evidence, but strong conclusions on food-related density dependence were constrained by the paucity of food availability data over the large spatial scales of this region. Our study highlights the importance of incorporating nonconstant conditions of species-environment relationships into predictive models of species distributions and population dynamics, and provides guidance for improved monitoring of fisheries and climate change impacts in the Southern Ocean.

High inter- and intraspecific niche overlap among three sympatrically breeding, closely related seabird species: Generalist foraging as an adaptation to a highly variable environment?

Ecological niche theory predicts sympatric species to show segregation in their spatio-temporal habitat utilization or diet as a strategy to avoid competition. Similarly, within species individuals may specialize on specific dietary resources or foraging habitats. Such individual specialization seems to occur particularly in environments with predictable resource distribution and limited environmental variability. Still, little is known about how seasonal environmental variability affects segregation of resources within species and between closely related sympatric species. The aim of the study was to investigate the foraging behaviour of three closely related and sympatrically breeding fulmarine petrels (Antarctic petrels Thalassoica antarctica, cape petrels Daption capense and southern fulmars Fulmarus glacialoides) in a seasonally highly variable environment (Prydz Bay, Antarctica) with the aim of assessing inter- and intraspecific overlap in utilized habitat, timing of foraging and diet and to identify foraging habitat preferences. We used GPS loggers with wet/dry sensors to assess spatial habitat utilization over the entire breeding season. Trophic overlap was investigated using stable isotope analysis based on blood, feathers and egg membranes. Foraging locations were identified using wet/dry data recorded by the GPS loggers and expectation-maximization binary clustering. Foraging habitat preferences were modelled using generalized additive models and model cross-validation. During incubation and chick-rearing, the utilization distribution of all three species overlapped significantly and species also overlapped in the timing of foraging during the day-partly during incubation and completely during chick-rearing. Isotopic centroids showed no significant segregation between at least two species for feathers and egg membranes, and among all species during incubation (reflected by blood). Within species, there was no individual specialization in foraging sites or environmental space. Furthermore, no single environmental covariate predicted foraging activity along trip trajectories. Instead, best-explanatory environmental covariates varied within and between individuals even across short temporal scales, reflecting a highly generalist behaviour of birds. Our results may be explained by optimal foraging theory. In the highly productive but spatio-temporally variable Antarctic environment, being a generalist may be key to finding mobile prey-even though this increases the potential for competition within and among sympatric species.

Author Correction: Time-lapse imagery and volunteer classifications from the Zooniverse Penguin Watch project.

An amendment to this paper has been published and can be accessed via a link at the top of the paper.

Nonbreeder birds at colonies display qualitatively similar seasonal mass change patterns as breeders.

The difficulty in studying nonbreeding birds means that little is known about them or their resource requirements, despite forming a large and significant component of a population. One way to assess food requirements is to examine changes in body mass, because it indicates the amount of food acquired. In terms of body mass changes, our expectation is that nonbreeders will either (a) be in poorer condition than the breeders which potentially explains why they do not breed or (b) remain at a stable higher mass as they are unconstrained by the physiological costs associated with rearing chicks. Here, we interrogate body mass datasets of breeding and nonbreeding birds of two penguin species to assess these predictions and determine whether differences in mass exist between these two groups throughout the breeding season. The first dataset is from a wild Adélie penguin population, where bird mass was recorded automatically and breeding status determined from a resighting program. A second population of captive gentoo penguins were weighed regularly each breeding season. We demonstrate that although there were times in each year when breeders were heavier than their nonbreeding counterparts for both populations, the mass changes showed qualitatively similar patterns throughout the breeding season irrespective of breeding status. Heavier breeders at times during the breeding season are not unexpected but the overall similar pattern of mass change irrespective of breeding status is in contrast to expectations. It appears that breeding status per se and the constraints that breeding places on birds are not the only driver of changes in mass throughout the breeding season and, although not explicitly studied here, the role of hormones in driving changes in appetite could be key to explain these results. These results present a significant step toward understanding food requirements of nonbreeders in avian populations.

Comparative population genomics reveals key barriers to dispersal in Southern Ocean penguins.

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 genomewide 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.

Time-lapse imagery and volunteer classifications from the Zooniverse Penguin Watch project.

Automated time-lapse cameras can facilitate reliable and consistent monitoring of wild animal populations. In this report, data from 73,802 images taken by 15 different Penguin Watch cameras are presented, capturing the dynamics of penguin (Spheniscidae; Pygoscelis spp.) breeding colonies across the Antarctic Peninsula, South Shetland Islands and South Georgia (03/2012 to 01/2014). Citizen science provides a means by which large and otherwise intractable photographic data sets can be processed, and here we describe the methodology associated with the Zooniverse project Penguin Watch, and provide validation of the method. We present anonymised volunteer classifications for the 73,802 images, alongside the associated metadata (including date/time and temperature information). In addition to the benefits for ecological monitoring, such as easy detection of animal attendance patterns, this type of annotated time-lapse imagery can be employed as a training tool for machine learning algorithms to automate data extraction, and we encourage the use of this data set for computer vision development.

Time-lapse imagery of Adélie penguins reveals differential winter strategies and breeding site occupation.

Polar seabirds adopt different over-wintering strategies to survive and build condition during the critical winter period. Penguin species either reside at the colony during the winter months or migrate long distances. Tracking studies and survey methods have revealed differences in winter migration routes among penguin species and colonies, dependent on both biotic and abiotic factors present. However, scan sampling methods are rarely used to reveal non-breeding behaviors during winter and little is known about presence at the colony site over this period. Here we show that Adélie penguins on the Yalour Islands in the Western Antarctic Peninsula (WAP) are present year-round at the colony and undergo a mid-winter peak in abundance during winter. We found a negative relationship between daylight hours and penguin abundance when either open water or compact ice conditions were present, suggesting that penguins return to the breeding colony when visibility is lowest for at-sea foraging and when either extreme low or high levels of sea ice exist offshore. In contrast, Adélie penguins breeding in East Antarctica were not observed at the colonies during winter, suggesting that Adélie penguins undergo differential winter strategies in the marginal ice zone on the WAP compared to those in East Antarctica. These results demonstrate that cameras can successfully monitor wildlife year-round in areas that are largely inaccessible during winter.

Circumpolar analysis of the Adélie Penguin reveals the importance of environmental variability in phenological mismatch.

Evidence of climate-change-driven shifts in plant and animal phenology have raised concerns that certain trophic interactions may be increasingly mismatched in time, resulting in declines in reproductive success. Given the constraints imposed by extreme seasonality at high latitudes and the rapid shifts in phenology seen in the Arctic, we would also expect Antarctic species to be highly vulnerable to climate-change-driven phenological mismatches with their environment. However, few studies have assessed the impacts of phenological change in Antarctica. Using the largest database of phytoplankton phenology, sea-ice phenology, and Adélie Penguin breeding phenology and breeding success assembled to date, we find that, while a temporal match between Penguin breeding phenology and optimal environmental conditions sets an upper limit on breeding success, only a weak relationship to the mean exists. Despite previous work suggesting that divergent trends in Adélie Penguin breeding phenology are apparent across the Antarctic continent, we find no such trends. Furthermore, we find no trend in the magnitude of phenological mismatch, suggesting that mismatch is driven by interannual variability in environmental conditions rather than climate-change-driven trends, as observed in other systems. We propose several criteria necessary for a species to experience a strong climate-change-driven phenological mismatch, of which several may be violated by this system.

Simultaneous DNA-based diet analysis of breeding, non-breeding and chick Adélie penguins.

As central place foragers, breeding penguins are restricted in foraging range by the need to return to the colony to feed chicks. Furthermore, breeding birds must balance energetic gain from self-feeding with the costs of returning to provision young. Non-breeding birds, however, are likely to be less restricted in foraging range and lack the high energy demands of provisioning, therefore may consume different prey to breeders. We used DNA dietary analysis to determine whether there was a difference in provisioning and self-feeding diet by identifying prey DNA in scat samples from breeding and chick Adélie penguins at two locations in East Antarctica. We also investigated diet differences between breeders and non-breeders at one site. Although previous work shows changing foraging behaviour between chick provisioning and self-feeding, our results suggest no significant differences in the main prey groups consumed by chicks and breeders at either site or between breeding stages. This may reflect the inability of penguins to selectively forage when provisioning, or resources were sufficient for all foraging needs. Conversely, non-breeders were found to consume different prey groups to breeders, which may reflect less restricted foraging ranges, breeders actively selecting particular prey during breeding or reduced foraging experience of non-breeders.

Proliferation of East Antarctic Adélie penguins in response to historical deglaciation.

BACKGROUND: Major, long-term environmental changes are projected in the Southern Ocean and these are likely to have impacts for marine predators such as the Adélie penguin (Pygoscelis adeliae). Decadal monitoring studies have provided insight into the short-term environmental sensitivities of Adélie penguin populations, particularly to sea ice changes. However, given the long-term nature of projected climate change, it is also prudent to consider the responses of populations to environmental change over longer time scales. We investigated the population trajectory of Adélie penguins during the last glacial-interglacial transition to determine how the species was affected by climate warming over millennia. We focussed our study on East Antarctica, which is home to 30 % of the global population of Adélie penguins. METHODS: Using mitochondrial DNA from extant colonies, we reconstructed the population trend of Adélie penguins in East Antarctica over the past 22,000 years using an extended Bayesian skyline plot method. To determine the relationship of East Antarctic Adélie penguins with populations elsewhere in Antarctica we constructed a phylogeny using mitochondrial DNA sequences. RESULTS: We found that the Adélie penguin population expanded 135-fold from approximately 14,000 years ago. The population growth was coincident with deglaciation in East Antarctica and, therefore, an increase in ice-free ground suitable for Adélie penguin nesting. Our phylogenetic analysis indicated that East Antarctic Adélie penguins share a common ancestor with Adélie penguins from the Antarctic Peninsula and Scotia Arc, with an estimated age of 29,000 years ago, in the midst of the last glacial period. This finding suggests that extant colonies in East Antarctica, the Scotia Arc and the Antarctic Peninsula were founded from a single glacial refuge. CONCLUSIONS: While changes in sea ice conditions are a critical driver of Adélie penguin population success over decadal and yearly timescales, deglaciation appears to have been the key driver of population change over millennia. This suggests that environmental drivers of population trends over thousands of years may differ to drivers over years or decades, highlighting the need to consider millennial-scale trends alongside contemporary data for the forecasting of species' abundance and distribution changes under future climate change scenarios.

Spatially Extensive Standardized Surveys Reveal Widespread, Multi-Decadal Increase in East Antarctic Adélie Penguin Populations.

Seabirds are considered to be useful and practical indicators of the state of marine ecosystems because they integrate across changes in the lower trophic levels and the physical environment. Signals from this key group of species can indicate broad scale impacts or response to environmental change. Recent studies of penguin populations, the most commonly abundant Antarctic seabirds in the west Antarctic Peninsula and western Ross Sea, have demonstrated that physical changes in Antarctic marine environments have profound effects on biota at high trophic levels. Large populations of the circumpolar-breeding Adélie penguin occur in East Antarctica, but direct, standardized population data across much of this vast coastline have been more limited than in other Antarctic regions. We combine extensive new population survey data, new population estimation methods, and re-interpreted historical survey data to assess decadal-scale change in East Antarctic Adélie penguin breeding populations. We show that, in contrast to the west Antarctic Peninsula and western Ross Sea where breeding populations have decreased or shown variable trends over the last 30 years, East Antarctic regional populations have almost doubled in abundance since the 1980's and have been increasing since the earliest counts in the 1960's. The population changes are associated with five-year lagged changes in the physical environment, suggesting that the changing environment impacts primarily on the pre-breeding age classes. East Antarctic marine ecosystems have been subject to a number of changes over the last 50 years which may have influenced Adélie penguin population growth, including decadal-scale climate variation, an inferred mid-20th century sea-ice contraction, and early-to-mid 20th century exploitation of fish and whale populations.

Re-constructing historical Adélie penguin abundance estimates by retrospectively accounting for detection bias.

Seabirds and other land-breeding marine predators are considered to be useful and practical indicators of the state of marine ecosystems because of their dependence on marine prey and the accessibility of their populations at breeding colonies. Historical counts of breeding populations of these higher-order marine predators are one of few data sources available for inferring past change in marine ecosystems. However, historical abundance estimates derived from these population counts may be subject to unrecognised bias and uncertainty because of variable attendance of birds at breeding colonies and variable timing of past population surveys. We retrospectively accounted for detection bias in historical abundance estimates of the colonial, land-breeding Adélie penguin through an analysis of 222 historical abundance estimates from 81 breeding sites in east Antarctica. The published abundance estimates were de-constructed to retrieve the raw count data and then re-constructed by applying contemporary adjustment factors obtained from remotely operating time-lapse cameras. The re-construction process incorporated spatial and temporal variation in phenology and attendance by using data from cameras deployed at multiple sites over multiple years and propagating this uncertainty through to the final revised abundance estimates. Our re-constructed abundance estimates were consistently higher and more uncertain than published estimates. The re-constructed estimates alter the conclusions reached for some sites in east Antarctica in recent assessments of long-term Adélie penguin population change. Our approach is applicable to abundance data for a wide range of colonial, land-breeding marine species including other penguin species, flying seabirds and marine mammals.

Climate change and Southern Ocean ecosystems I: how changes in physical habitats directly affect marine biota.

Antarctic and Southern Ocean (ASO) marine ecosystems have been changing for at least the last 30 years, including in response to increasing ocean temperatures and changes in the extent and seasonality of sea ice; the magnitude and direction of these changes differ between regions around Antarctica that could see populations of the same species changing differently in different regions. This article reviews current and expected changes in ASO physical habitats in response to climate change. It then reviews how these changes may impact the autecology of marine biota of this polar region: microbes, zooplankton, salps, Antarctic krill, fish, cephalopods, marine mammals, seabirds, and benthos. The general prognosis for ASO marine habitats is for an overall warming and freshening, strengthening of westerly winds, with a potential pole-ward movement of those winds and the frontal systems, and an increase in ocean eddy activity. Many habitat parameters will have regionally specific changes, particularly relating to sea ice characteristics and seasonal dynamics. Lower trophic levels are expected to move south as the ocean conditions in which they are currently found move pole-ward. For Antarctic krill and finfish, the latitudinal breadth of their range will depend on their tolerance of warming oceans and changes to productivity. Ocean acidification is a concern not only for calcifying organisms but also for crustaceans such as Antarctic krill; it is also likely to be the most important change in benthic habitats over the coming century. For marine mammals and birds, the expected changes primarily relate to their flexibility in moving to alternative locations for food and the energetic cost of longer or more complex foraging trips for those that are bound to breeding colonies. Few species are sufficiently well studied to make comprehensive species-specific vulnerability assessments possible. Priorities for future work are discussed.

Adélie penguin population diet monitoring by analysis of food DNA in scats.

The Adélie penguin is the most important animal currently used for ecosystem monitoring in the Southern Ocean. The diet of this species is generally studied by visual analysis of stomach contents; or ratios of isotopes of carbon and nitrogen incorporated into the penguin from its food. There are significant limitations to the information that can be gained from these methods. We evaluated population diet assessment by analysis of food DNA in scats as an alternative method for ecosystem monitoring with Adélie penguins as an indicator species. Scats were collected at four locations, three phases of the breeding cycle, and in four different years. A novel molecular diet assay and bioinformatics pipeline based on nuclear small subunit ribosomal RNA gene (SSU rDNA) sequencing was used to identify prey DNA in 389 scats. Analysis of the twelve population sample sets identified spatial and temporal dietary change in Adélie penguin population diet. Prey diversity was found to be greater than previously thought. Krill, fish, copepods and amphipods were the most important food groups, in general agreement with other Adélie penguin dietary studies based on hard part or stable isotope analysis. However, our DNA analysis estimated that a substantial portion of the diet was gelatinous groups such as jellyfish and comb jellies. A range of other prey not previously identified in the diet of this species were also discovered. The diverse prey identified by this DNA-based scat analysis confirms that the generalist feeding of Adélie penguins makes them a useful indicator species for prey community composition in the coastal zone of the Southern Ocean. Scat collection is a simple and non-invasive field sampling method that allows DNA-based estimation of prey community differences at many temporal and spatial scales and provides significant advantages over alternative diet analysis approaches.