Tracking of marine predators to protect Southern Ocean ecosystems.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change1,2. Mitigation requires the identification and protection of Areas of Ecological Significance (AESs), which have so far not been determined at the ocean-basin scale. Here, using assemblage-level tracking of marine predators, we identify AESs for this globally important region and assess current threats and protection levels. Integration of more than 4,000 tracks from 17 bird and mammal species reveals AESs around sub-Antarctic islands in the Atlantic and Indian Oceans and over the Antarctic continental shelf. Fishing pressure is disproportionately concentrated inside AESs, and climate change over the next century is predicted to impose pressure on these areas, particularly around the Antarctic continent. At present, 7.1% of the ocean south of 40°S is under formal protection, including 29% of the total AESs. The establishment and regular revision of networks of protection that encompass AESs are needed to provide long-term mitigation of growing pressures on Southern Ocean ecosystems.

The effect of climate change on avian offspring production: A global meta-analysis.

Climate change affects timing of reproduction in many bird species, but few studies have investigated its influence on annual reproductive output. Here, we assess changes in the annual production of young by female breeders in 201 populations of 104 bird species (N = 745,962 clutches) covering all continents between 1970 and 2019. Overall, average offspring production has declined in recent decades, but considerable differences were found among species and populations. A total of 56.7% of populations showed a declining trend in offspring production (significant in 17.4%), whereas 43.3% exhibited an increase (significant in 10.4%). The results show that climatic changes affect offspring production through compounded effects on ecological and life history traits of species. Migratory and larger-bodied species experienced reduced offspring production with increasing temperatures during the chick-rearing period, whereas smaller-bodied, sedentary species tended to produce more offspring. Likewise, multi-brooded species showed increased breeding success with increasing temperatures, whereas rising temperatures were unrelated to reproductive success in single-brooded species. Our study suggests that rapid declines in size of bird populations reported by many studies from different parts of the world are driven only to a small degree by changes in the production of young.

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.

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.

Ageing simulation in health and social care education: A mixed methods systematic review.

AIM: To identify, evaluate and summarize evidence from qualitative, quantitative and mixed method studies conducted using age suits or other age simulation equipment, with health and social care students. DESIGN: Convergent segregated mixed method review design as outlined by the Johanna Briggs Institute. DATA SOURCES: CINAHL (+ with Full Text), MEDLINE, PsycINFO, PubMed, SocINDEX, Web of Science, Cochrane Library, Emerald Insight, Proquest nursing, Science Direct, Wiley Online and BioMed Central (January 2000-January 2020). REVIEW METHODS: Convergent segregated synthesis was used to synthesize evidence from the studies and the MERSQI checklist used to appraise quality. RESULTS: A total of 23 studies were reviewed: one randomized control, two post-test only randomized control, three quasi-experimental, 15 one-group pre/post studies and two qualitative studies. Of the seventeen studies carrying out inferential statistics on attitude scores post intervention, 11 reported an improvement, three indicated no significant change and three reported worsening scores. Key themes included use of appropriate scales, type of equipment used, location and length of interactions, debriefing and contextualization of interventions in broader teaching. CONCLUSION: The impact of ageing simulation interventions on health and social care student's attitudes to older people was predominantly positive. However, further high-quality research is warranted to understand the optimal use of such interventions in the context of health care for a growing ageing population. IMPACT: It is important health and social care staff have appropriate knowledge and training to enable them to provide high-quality care to older people and challenge potential ageism in the system. This review adds to the body of work around the use of simulation and experiential learning to educate health and social care students about ageing and ageism. It also offers recommendations for using ageing simulations effectively to inform attitudes of prospective professionals who will influence future health and social care.

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.

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.

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.

The influence of historical climate changes on Southern Ocean marine predator populations: a comparative analysis.

The Southern Ocean ecosystem is undergoing rapid physical and biological changes that are likely to have profound implications for higher-order predators. Here, we compare the long-term, historical responses of Southern Ocean predators to climate change. We examine palaeoecological evidence for changes in the abundance and distribution of seabirds and marine mammals, and place these into context with palaeoclimate records in order to identify key environmental drivers associated with population changes. Our synthesis revealed two key factors underlying Southern Ocean predator population changes; (i) the availability of ice-free ground for breeding and (ii) access to productive foraging grounds. The processes of glaciation and sea ice fluctuation were key; the distributions and abundances of elephant seals, snow petrels, gentoo, chinstrap and Adélie penguins all responded strongly to the emergence of new breeding habitat coincident with deglaciation and reductions in sea ice. Access to productive foraging grounds was another limiting factor, with snow petrels, king and emperor penguins all affected by reduced prey availability in the past. Several species were isolated in glacial refugia and there is evidence that refuge populations were supported by polynyas. While the underlying drivers of population change were similar across most Southern Ocean predators, the individual responses of species to environmental change varied because of species specific factors such as dispersal ability and environmental sensitivity. Such interspecific differences are likely to affect the future climate change responses of Southern Ocean marine predators and should be considered in conservation plans. Comparative palaeoecological studies are a valuable source of long-term data on species' responses to environmental change that can provide important insights into future climate change responses. This synthesis highlights the importance of protecting productive foraging grounds proximate to breeding locations, as well as the potential role of polynyas as future Southern Ocean refugia.

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.

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.

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.

Changes in seed dispersal processes and the potential for between-patch connectivity for an arid land daisy.

Dispersal is a major and critical process in population biology that has been particularly challenging to study. Animals can have major roles in seed dispersal even in species that do not appear specifically adapted to animal-aided dispersal. This can occur by two processes: direct movement of diaspores by animals and modification of landscape characteristics by animals in ways that greatly influence dispersal. We exploited the production of large, persistent dispersal structures (seed heads, henceforth) by Erodiophyllum elderi (Asteraceae), a daisy from arid Australia, to further understand secondary dispersal. Seed head dispersal on and off animal tracks in eight E. elderi patches was monitored for 9.5 months by periodically recording the location of marked seed heads. Sites were located inside a reserve that excludes sheep but not kangaroos, and in a nearby area with both kangaroos and sheep. The distance moved and likelihood of seed head movement was higher in areas with sheep, and especially along animal tracks. There was clear evidence that seed heads were channeled down animal tracks during large rainfall events. Seed head dispersal away from patches occurred to a limited extent via their physical contact with sheep and potentially via wind dispersal. Thus, the advantages of this study system allowed us to demonstrate the two postulated effects of herbivores on dispersal via direct movement of seed heads, and two distinct indirect effects through landscape modification by herbivores from the creation of animal tracks and the denudation of vegetation.

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.

New and legacy persistent organic pollutants (POPs) in breeding seabirds from the East Antarctic.

Persistent organic pollutants (POPs) are pervasive and a significant threat to the environment worldwide. Yet, reports of POP levels in Antarctic seabirds based on blood are scarce, resulting in significant geographical gaps. Blood concentrations offer a snapshot of contamination within live populations, and have been used widely for Arctic and Northern Hemisphere seabird species but less so in Antarctica. This paper presents levels of legacy POPs (polychlorinated biphenyls (PCBs), organochlorine pesticides (OCPs) and polybrominated diphenyl ethers (PBDEs)) and novel brominated flame retardants (NBFRs) in the blood of five Antarctic seabird species breeding within Prydz Bay, East Antarctica. Legacy PCBs and OCPs were detected in all species sampled, with Adélie penguins showing comparatively high ∑PCB levels (61.1 ± 87.6 ng/g wet weight (ww)) compared to the four species of flying seabirds except the snow petrel (22.5 ± 15.5 ng/g ww), highlighting that legacy POPs are still present within Antarctic wildlife despite decades-long bans. Both PBDEs and NBFRs were detected in trace levels for all species and hexabromobenzene (HBB) was quantified in cape petrels (0.3 ± 0.2 ng/g ww) and snow petrels (0.2 ± 0.1 ng/g ww), comparable to concentrations found in Arctic seabirds. These results fill a significant data gap within the Antarctic region for POPs studies, representing a crucial step forward assessing the fate and impact of legacy POPs contamination in the Antarctic environment.

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.

Adélie penguin survival: age structure, temporal variability and environmental influences.

The driving factors of survival, a key demographic process, have been particularly challenging to study, especially for winter migratory species such as the Adélie penguin (Pygoscelis adeliae). While winter environmental conditions clearly influence Antarctic seabird survival, it has been unclear to which environmental features they are most likely to respond. Here, we examine the influence of environmental fluctuations, broad climatic conditions and the success of the breeding season prior to winter on annual survival of an Adélie penguin population using mark-recapture models based on penguin tag and resight data over a 16-year period. This analysis required an extension to the basic Cormack-Jolly-Seber model by incorporating age structure in recapture and survival sub-models. By including model covariates, we show that survival of older penguins is primarily related to the amount and concentration of ice present in their winter foraging grounds. In contrast, fledgling and yearling survival depended on other factors in addition to the physical marine environment and outcomes of the previous breeding season, but we were unable to determine what these were. The relationship between sea-ice and survival differed with penguin age: extensive ice during the return journey to breeding colonies was detrimental to survival for the younger penguins, whereas either too little or too much ice (between 15 and 80% cover) in the winter foraging grounds was detrimental for adults. Our results demonstrate that predictions of Adélie penguin survival can be improved by taking into account penguin age, prior breeding conditions and environmental features.

Horizontal-vertical movement relationships: Adélie penguins forage continuously throughout provisioning trips.

BACKGROUND: Diving marine predators forage in a three-dimensional environment, adjusting their horizontal and vertical movement behaviour in response to environmental conditions and the spatial distribution of prey. Expectations regarding horizontal-vertical movements are derived from optimal foraging theories, however, inconsistent empirical findings across a range of taxa suggests these behavioural assumptions are not universally applicable. METHODS: Here, we examined how changes in horizontal movement trajectories corresponded with diving behaviour and marine environmental conditions for a ubiquitous Southern Ocean predator, the Adélie penguin. Integrating extensive telemetry-based movement and environmental datasets for chick-rearing Adélie penguins at Béchervaise Island, we tested the relationships between horizontal move persistence (continuous scale indicating low ['resident'] to high ['directed'] movement autocorrelation), vertical dive effort and environmental variables. RESULTS: Penguins dived continuously over the course of their foraging trips and lower horizontal move persistence corresponded with less intense foraging activity, likely indicative of resting behaviour. This challenges the traditional interpretation of horizontal-vertical movement relationships based on optimal foraging models, which assumes increased residency within an area translates to increased foraging activity. Movement was also influenced by different environmental conditions during the two stages of chick-rearing: guard and crèche. These differences highlight the strong seasonality of foraging habitat for chick-rearing Adélie penguins at Béchervaise Island. CONCLUSIONS: Our findings advance our understanding of the foraging behaviour for this marine predator and demonstrates the importance of integrating spatial location and behavioural data before inferring habitat use.