Incorporating mesopelagic fish into the evaluation of conservation areas for marine living resources under climate change scenarios.

Mesopelagic fish (meso-fish) are central species within the Southern Ocean (SO). However, their ecosystem role and adaptive capacity to climate change are rarely integrated into protected areas assessments. This is a pity given their importance as crucial prey and predators in food webs, coupled with the impacts of climate change. Here, we estimate the habitat distribution of nine meso-fish using an ensemble model approach (MAXENT, random forest, and boosted regression tree). Four climate model simulations were used to project their distribution under two representative concentration pathways (RCP4.5 and RCP8.5) for short-term (2006-2055) and long-term (2050-2099) periods. In addition, we assess the ecological representativeness of protected areas under climate change scenarios using meso-fish as indicator species. Our models show that all species shift poleward in the future. Lanternfishes (family Myctophidae) are predicted to migrate poleward more than other families (Paralepididae, Nototheniidae, Bathylagidae, and Gonostomatidae). In comparison, lanternfishes were projected to increase habitat area in the eastern SO but lose area in the western SO; the opposite was projected for species in other families. Important areas (IAs) of meso-fish are mainly distributed near the Antarctic Peninsula and East Antarctica. Negotiated protected area cover 23% of IAs at present and 38% of IAs in the future (RCP8.5, long-term future). Many IAs of meso-fish still need to be included in protected areas, such as the Prydz Bay and the seas around the Antarctic Peninsula. Our results provide a framework for evaluating protected areas incorporating climate change adaptation strategies for protected areas management. Supplementary Information: The online version contains supplementary material available at 10.1007/s42995-023-00188-9.

The benthic-pelagic continuum: Age class and sex differences in the use of the vertical dimension by a rare pinniped.

Sea lions as a group, display strong site fidelity, and varying degrees of vulnerability to environmental change, disease and fisheries interactions. One of the rarest pinnipeds, the New Zealand sea lion (NZSL, Phocarctos hookeri) has a very restricted breeding range. At Campbell Island/Motu Ihupuku, one of the two primary breeding sites, at-sea foraging behaviour is unknown. We hypothesised that NZSL of various sex and age classes would utilise the water column differently due to differing physiological constraints and therefore have different accessibility to prey resources. We tested whether sea lion diving behaviour varied in relation to (i) age and sex class, (ii) time of day and (iii) water depth. We also hypothesised that the proportion of benthic/pelagic diving, and consequently risk of fisheries interaction, would vary in relation to age and sex. Satellite telemetry tags were deployed on 25 NZSL from a range of age/sex classes recording dive depth, duration and location. Adult females and juveniles used inshore, benthic habitats, while sub-adult males also utilised benthic habitats, they predominantly used pelagic habitat at greater distances from the island. Adult females and juveniles exhibited shorter dives than the same age/sex classes at the Auckland Islands, suggesting a lower dive effort for these age/sex classes at Campbell Island/Motu Ihupuku. Adult females dived more frequently than other age/sex classes, likely operating closer to their physiological limits; however, further data for this age class is needed. Sub-adult male use of pelagic prey may increase their exposure to mid-water trawls; however, further research detailing the degree of spatial overlap with fisheries is required. This study highlights the utility of spatially explicit dive data to predict vertical habitat use, niche separation of various age and sex classes of marine predators and attribute potential fisheries interaction risk in relation to predator habitat use.

A vision for incorporating human mobility in the study of human-wildlife interactions.

As human activities increasingly shape land- and seascapes, understanding human-wildlife interactions is imperative for preserving biodiversity. Habitats are impacted not only by static modifications, such as roads, buildings and other infrastructure, but also by the dynamic movement of people and their vehicles occurring over shorter time scales. Although there is increasing realization that both components of human activity substantially affect wildlife, capturing more dynamic processes in ecological studies has proved challenging. Here we propose a conceptual framework for developing a 'dynamic human footprint' that explicitly incorporates human mobility, providing a key link between anthropogenic stressors and ecological impacts across spatiotemporal scales. Specifically, the dynamic human footprint integrates a range of metrics to fully acknowledge the time-varying nature of human activities and to enable scale-appropriate assessments of their impacts on wildlife behaviour, demography and distributions. We review existing terrestrial and marine human-mobility data products and provide a roadmap for how these could be integrated and extended to enable more comprehensive analyses of human impacts on biodiversity in the Anthropocene.

Environmental Drivers of Growth and Oxidative Status during Early Life in a Long-Lived Antarctic Seabird, the Adélie Penguin.

AbstractIn vertebrates, developmental conditions can have long-term effects on individual performance. It is increasingly recognized that oxidative stress could be one physiological mechanism connecting early-life experience to adult phenotype. Accordingly, markers of oxidative status could be useful for assessing the developmental constraints encountered by offspring. Although some studies have demonstrated that developmental constraints are associated with high levels of oxidative stress in offspring, it remains unclear how growth, parental behavior, and brood competition may altogether affect oxidative stress in long-lived species in the wild. Here, we investigated this question in a long-lived Antarctic bird species by testing the impact of brood competition (e.g., brood size and hatching order) on body mass and on two markers of oxidative damage in Adélie penguin chicks. We also examined the influence of parental effort (i.e., foraging trip duration) and parental body condition on chick body mass and oxidative damage. First, we found that brood competition and parental traits had significant impacts on chick body mass. Second, we found that chick age and, to a lesser extent, chick body mass were two strong determinants of the levels of oxidative damage in Adélie penguin chicks. Finally, and importantly, we also found that brood competition significantly increased the levels of one marker of oxidative damage and was associated with a lower survival probability. However, parental effort and parental condition were not significantly linked to chick levels of oxidative damage. Overall, our study demonstrates that sibling competition can generate an oxidative cost even for this long-lived Antarctic species with a limited brood size (maximum of two chicks).

Extreme polygyny results in intersex differences in age-dependent survival of a highly dimorphic marine mammal.

Developmental differences in vital rates are especially profound in polygamous mating systems. Southern elephant seals (Mirounga leonina) are highly dimorphic and extremely polygynous marine mammals. A demographic model, supported by long-term capture-mark-recapture records, investigated the influence of sex and age on survival in this species. The study revealed clear differences between female and male age-dependent survival rates. Overall juvenile survival estimates were stable around 80-85% for both sexes. However, male survival estimates were 5-10% lower than females in the same age classes until 8 years of age. At this point, male survival decreased rapidly to 50% ± 10% while female estimates remained constant at 80% ± 5%. Different energetic requirements could underpin intersex differences in adult survival. However, the species' strong sexual dimorphism diverges during early juvenile development when sex-specific survival rates were less distinct. Maximizing growth is especially advantageous for males, with size being a major determinant of breeding probability. Maturing males may employ a high-risk high-reward foraging strategy to compensate for extensive sexual selection pressures and sex-specific energetic needs. Our findings suggest sex-specific adult survival is a result of in situ ecological interactions and evolutionary specialization associated with being a highly polygynous marine predator.

Long-term stability in the circumpolar foraging range of a Southern Ocean predator between the eras of whaling and rapid climate change.

Assessing environmental changes in Southern Ocean ecosystems is difficult due to its remoteness and data sparsity. Monitoring marine predators that respond rapidly to environmental variation may enable us to track anthropogenic effects on ecosystems. Yet, many long-term datasets of marine predators are incomplete because they are spatially constrained and/or track ecosystems already modified by industrial fishing and whaling in the latter half of the 20th century. Here, we assess the contemporary offshore distribution of a wide-ranging marine predator, the southern right whale (SRW, Eubalaena australis), that forages on copepods and krill from ~30°S to the Antarctic ice edge (>60°S). We analyzed carbon and nitrogen isotope values of 1,002 skin samples from six genetically distinct SRW populations using a customized assignment approach that accounts for temporal and spatial variation in the Southern Ocean phytoplankton isoscape. Over the past three decades, SRWs increased their use of mid-latitude foraging grounds in the south Atlantic and southwest (SW) Indian oceans in the late austral summer and autumn and slightly increased their use of high-latitude (>60°S) foraging grounds in the SW Pacific, coincident with observed changes in prey distribution and abundance on a circumpolar scale. Comparing foraging assignments with whaling records since the 18th century showed remarkable stability in use of mid-latitude foraging areas. We attribute this consistency across four centuries to the physical stability of ocean fronts and resulting productivity in mid-latitude ecosystems of the Southern Ocean compared with polar regions that may be more influenced by recent climate change.

Potential for redistribution of post-moult habitat for Eudyptes penguins in the Southern Ocean under future climate conditions.

Anthropogenic climate change is resulting in spatial redistributions of many species. We assessed the potential effects of climate change on an abundant and widely distributed group of diving birds, Eudyptes penguins, which are the main avian consumers in the Southern Ocean in terms of biomass consumption. Despite their abundance, several of these species have undergone population declines over the past century, potentially due to changing oceanography and prey availability over the important winter months. We used light-based geolocation tracking data for 485 individuals deployed between 2006 and 2020 across 10 of the major breeding locations for five taxa of Eudyptes penguins. We used boosted regression tree modelling to quantify post-moult habitat preference for southern rockhopper (E. chrysocome), eastern rockhopper (E. filholi), northern rockhopper (E. moseleyi) and macaroni/royal (E. chrysolophus and E. schlegeli) penguins. We then modelled their redistribution under two climate change scenarios, representative concentration pathways RCP4.5 and RCP8.5 (for the end of the century, 2071-2100). As climate forcings differ regionally, we quantified redistribution in the Atlantic, Central Indian, East Indian, West Pacific and East Pacific regions. We found sea surface temperature and sea surface height to be the most important predictors of current habitat for these penguins; physical features that are changing rapidly in the Southern Ocean. Our results indicated that the less severe RCP4.5 would lead to less habitat loss than the more severe RCP8.5. The five taxa of penguin may experience a general poleward redistribution of their preferred habitat, but with contrasting effects in the (i) change in total area of preferred habitat under climate change (ii) according to geographic region and (iii) the species (macaroni/royal vs. rockhopper populations). Our results provide further understanding on the regional impacts and vulnerability of species to climate change.

Myctobase, a circumpolar database of mesopelagic fishes for new insights into deep pelagic prey fields.

The global importance of mesopelagic fish is increasingly recognised, but they remain poorly studied. This is particularly true in the Southern Ocean, where mesopelagic fishes are both key predators and prey, but where the remote environment makes sampling challenging. Despite this, multiple national Antarctic research programs have undertaken regional sampling of mesopelagic fish over several decades. However, data are dispersed, and sampling methodologies often differ precluding comparisons and limiting synthetic analyses. We identified potential data holders by compiling a metadata catalogue of existing survey data for Southern Ocean mesopelagic fishes. Data holders contributed 17,491 occurrence and 11,190 abundance records from 4780 net hauls from 72 different research cruises. Data span across 37 years from 1991 to 2019 and include trait-based information (length, weight, maturity). The final dataset underwent quality control processes and detailed metadata was provided for each sampling event. This dataset can be accessed through Zenodo. Myctobase will enhance research capacity by providing the broadscale baseline data necessary for observing and modelling mesopelagic fishes.

The role of allochrony in influencing interspecific differences in foraging distribution during the non-breeding season between two congeneric crested penguin species.

Mechanisms promoting coexistence between closely related species are fundamental for maintaining species diversity. Mechanisms of niche differentiation include allochrony which offsets the peak timing of resource utilisation between species. Many studies focus on spatial and temporal niche partitioning during the breeding season, few have investigated the role allochrony plays in influencing interspecific segregation of foraging distribution and ecology between congeneric species during the non-breeding season. We investigated the non-breeding migrations of Snares (Eudyptes robustus) and Fiordland penguins (Eudyptes pachyrhynchus), closely related species breeding between 100-350 km apart whose migration phenology differs by two months. Using light geolocation tracking, we examined the degree of overlap given the observed allochrony and a hypothetical scenario where the species commence migration simultaneously. We found that Fiordland penguins migrated to the Sub-Antarctic Frontal Zone and Polar Frontal Zone in the austral autumn whereas Snares penguins disperse westwards staying north of the Sub-Tropical Front in the austral winter. Our results suggest that allochrony is likely to be at the root of segregation because the relative profitability of the different water masses that the penguins forage in changes seasonally which results in the two species utilising different areas over their core non-breeding periods. Furthermore, allochrony reduces relatively higher levels of spatiotemporal overlap during the departure and arrival periods, when the close proximity of the two species' colonies would cause the birds to congregate in similar areas, resulting in high interspecific competition just before the breeding season. Available evidence from other studies suggests that the shift in phenology between these species has arisen from adaptive radiation and phenological matching to the seasonality of local resource availability during the breeding season and reduced competitive overlap over the non-breeding season is likely to be an incidental outcome.

Sex, body size, and boldness shape the seasonal foraging habitat selection in southern elephant seals.

Selecting foraging habitat is a fundamental behavior in the life of organisms as it directly links resource acquisition to fitness. Differences in habitat selection among individuals may arise from several intrinsic and extrinsic factors, and yet, their interaction has been given little attention in the study of wild populations. We combine sex, body size, and boldness to explain individual differences in the seasonal foraging habitat selection of southern elephant seals (Mirounga leonina) from the Kerguelen Archipelago. We hypothesize that habitat selection is linked to the trade-off between resource acquisition and risk, and that individuals differ in their position along this trade-off because of differences in reproductive strategies, life stages, and metabolic requirements. Before the post-molt foraging trip, we used a novel object approach test to quantify the boldness of 28 subadult and adult females and 42 subadult males and equipped them with data loggers to track their movements at sea. Subadult males selected neritic and oceanic habitats, whereas females mostly selected less productive oceanic habitats. Both sexes showed a seasonal shift from Antarctic habitats in the south in the summer to the free of ice subantarctic and subtropical habitats in the north in the winter. Males avoided oceanic habitats and selected more productive neritic and Antarctic habitats with body size mostly in the winter. Bolder males selected northern warmer waters in winter, while shyer ones selected the Kerguelen plateau and southern colder oceanic waters. Bolder females selected the Kerguelen plateau in the summer when prey profitability is assumed to be the highest. This study not only provides new insights into the spatiotemporal foraging ecology of elephant seals in relation to personality but also emphasizes the relevance of combining several intrinsic and extrinsic factors in understanding among-individual variation in space use essential in wildlife management and conservation.

Elephant seal foraging success is enhanced in Antarctic coastal polynyas.

Antarctic polynyas are persistent open water areas which enable early and large seasonal phytoplankton blooms. This high primary productivity, boosted by iron supply from coastal glaciers, attracts organisms from all trophic levels to form a rich and diverse community. How the ecological benefit of polynya productivity is translated to the highest trophic levels remains poorly resolved. We studied 119 southern elephant seals feeding over the Antarctic shelf and demonstrated that: (i) 96% of seals foraging here used polynyas, with individuals spending on average 62% of their time there; (ii) the seals exhibited more area-restricted search behaviour when in polynyas; and (iii) these seals gained more energy (indicated by increased buoyancy from greater fat stores) when inside polynyas. This higher-quality foraging existed even when ice was not present in the study area, indicating that these are important and predictable foraging grounds year-round. Despite these energetic advantages from using polynyas, not all the seals used them extensively. Factors other than food supply may influence an individual's choice in their use of feeding grounds, such as exposure to predation or the probability of being able to return to distant sub-Antarctic breeding sites.

Climate change impacts on seabirds and marine mammals: The importance of study duration, thermal tolerance and generation time.

Understanding climate change impacts on top predators is fundamental to marine biodiversity conservation, due to their increasingly threatened populations and their importance in marine ecosystems. We conducted a systematic review of the effects of climate change (prolonged, directional change) and climate variability on seabirds and marine mammals. We extracted data from 484 studies (4808 published studies were reviewed), comprising 2215 observations on demography, phenology, distribution, diet, behaviour, body condition and physiology. The likelihood of concluding that climate change had an impact increased with study duration. However, the temporal thresholds for the effects of climate change to be discernibly varied from 10 to 29 years depending on the species, the biological response and the oceanic study region. Species with narrow thermal ranges and relatively long generation times were more often reported to be affected by climate change. This provides an important framework for future assessments, with guidance on response- and region-specific temporal dimensions that need to be considered when reporting effects of climate change. Finally, we found that tropical regions and non-breeding life stages were poorly covered in the literature, a concern that should be addressed to enable a better understanding of the vulnerability of marine predators to climate change.

The cost of a meal: factors influencing prey profitability in Australian fur seals.

Knowledge of the factors shaping the foraging behaviour of species is central to understanding their ecosystem role and predicting their response to environmental variability. To maximise survival and reproduction, foraging strategies must balance the costs and benefits related to energy needed to pursue, manipulate, and consume prey with the nutritional reward obtained. While such information is vital for understanding how changes in prey assemblages may affect predators, determining these components is inherently difficult in cryptic predators. The present study used animal-borne video data loggers to investigate the costs and benefits related to different prey types for female Australian fur seals (Arctocephalus pusillus doriferus), a primarily benthic foraging species in the low productivity Bass Strait, south-eastern Australia. A total of 1,263 prey captures, resulting from 2,027 prey detections, were observed in 84.5 h of video recordings from 23 individuals. Substantial differences in prey pursuit and handling times, gross energy gain and total energy expenditure were observed between prey types. Importantly, the profitability of prey was not significantly different between prey types, with the exception of elasmobranchs. This study highlights the benefit of animal-borne video data loggers for understanding the factors that influence foraging decisions in predators. Further studies incorporating search times for different prey types would further elucidate how profitability differs with prey type.

Influence of environmental variation on spatial distribution and habitat-use in a benthic foraging marine predator.

The highly dynamic nature of the marine environment can have a substantial influence on the foraging behaviour and spatial distribution of marine predators, particularly in pelagic marine systems. However, knowledge of the susceptibility of benthic marine predators to environmental variability is limited. This study investigated the influence of local-scale environmental conditions and large-scale climate indices on the spatial distribution and habitat use in the benthic foraging Australian fur seal (Arctocephalus pusillus doriferus; AUFS). Female AUFS provisioning pups were instrumented with GPS or ARGOS platform terminal transmitter tags during the austral winters of 2001-2019 at Kanowna Island, south-eastern Australia. Individuals were most susceptible to changes in the Southern Oscillation Index that measures the strength of the El Niño Southern Oscillation, with larger foraging ranges, greater distances travelled and more dispersed movement associated with 1-yr lagged La Niña-like conditions. Additionally, the total distance travelled was negatively correlated with the current year sea surface temperature and 1-yr lagged Indian Ocean Dipole, and positively correlated with 1-yr lagged chlorophyll-a concentration. These results suggest that environmental variation may influence the spatial distribution and availability of prey, even within benthic marine systems.

A prediction and imputation method for marine animal movement data.

Data prediction and imputation are important parts of marine animal movement trajectory analysis as they can help researchers understand animal movement patterns and address missing data issues. Compared with traditional methods, deep learning methods can usually provide enhanced pattern extraction capabilities, but their applications in marine data analysis are still limited. In this research, we propose a composite deep learning model to improve the accuracy of marine animal trajectory prediction and imputation. The model extracts patterns from the trajectories with an encoder network and reconstructs the trajectories using these patterns with a decoder network. We use attention mechanisms to highlight certain extracted patterns as well for the decoder. We also feed these patterns into a second decoder for prediction and imputation. Therefore, our approach is a coupling of unsupervised learning with the encoder and the first decoder and supervised learning with the encoder and the second decoder. Experimental results demonstrate that our approach can reduce errors by at least 10% on average comparing with other methods.

Climate influences on female survival in a declining population of southern elephant seals (Mirounga leonina).

The Southern Ocean has been disproportionately affected by climate change and is therefore an ideal place to study the influence of changing environmental conditions on ecosystems. Changes in the demography of predator populations are indicators of broader shifts in food web structure, but long-term data are required to study these effects. Southern elephant seals (Mirounga leonina) from Macquarie Island have consistently decreased in population size while all other major populations across the Southern Ocean have recently stabilized or are increasing. Two long-term mark-recapture studies (1956-1967 and 1993-2009) have monitored this population, which provides an opportunity to investigate demographic performance over a range of climatic conditions. Using a 9-state matrix population model, we estimated climate influences on female survival by incorporating two major climatic indices into our model: The Southern Annular Mode (SAM) and the Southern Oscillation Index (SOI). Our best model included a 1 year lagged effect of SAM and an unlagged SOI as covariates. A positive relationship with SAM1 (lagged) related the previous year's SAM with juvenile survival, potentially due to changes in local prey availability surrounding Macquarie Island. The unlagged SOI had a negative effect on both juvenile and adult seals, indicating that sea ice dynamics and access to foraging grounds on the East Antarctic continental shelf could explain the different contributions of ENSO events on the survival of females in this population.

Intertrip consistency in hunting behavior improves foraging success and efficiency in a marine top predator.

Substantial variation in foraging strategies can exist within populations, even those typically regarded as generalists. Specializations arise from the consistent exploitation of a narrow behavioral, spatial or dietary niche over time, which may reduce intraspecific competition and influence adaptability to environmental change. However, few studies have investigated whether behavioral consistency confers benefits at the individual and/or population level. While still recovering from commercial sealing overexploitation, Australian fur seals (AUFS; Arctocephalus pusillus doriferus) represent the largest marine predator biomass in south-eastern Australia. During lactation, female AUFS adopt a central-place foraging strategy and are, thus, vulnerable to changes in prey availability. The present study investigated the population-level repeatability and individual consistency in foraging behavior of 34 lactating female AUFS at a south-east Australian breeding colony between 2006 and 2019. Additionally, the influence of individual-level behavioral consistency on indices of foraging success and efficiency during benthic diving was determined. Low to moderate population-level repeatability was observed across foraging behaviors, with the greatest repeatability in the mean bearing and modal dive depth. Individual-level consistency was greatest for the proportion of benthic diving, total distance travelled, and trip duration. Indices of benthic foraging success and efficiency were positively influenced by consistency in the proportion of benthic diving, trip duration and dive rate but not influenced by consistency in bearing to most distal point, dive depth or foraging site fidelity. The results of the present study provide evidence of the benefits of consistency for individuals, which may have flow-on effects at the population level.

Predicting the distribution of foraging seabirds during a period of heightened environmental variability.

Quantifying the links between the marine environment, prey occurrence, and predator distribution is the first step towards identifying areas of biological importance for marine spatial planning. Events such as marine heatwaves result in an anomalous change in the physical environment, which can lead to shifts in the structure, biomass, and distribution of lower trophic levels. As central-place foragers, seabirds are vulnerable to changes in their foraging grounds during the breeding season. We first quantified spatiotemporal variability in the occurrence and biomass of prey in response to an abrupt change in oceanography as a result of a marine heatwave event. Secondly, using multivariate techniques and machine learning, we investigated if differences in the foraging technique and prey of seabirds resulted in varying responses to changes in prey occurrence and the environment over a 2.5-yr period. We found that the main variables correlated with seabird distribution were also important in structuring the occurrence and biomass of prey; sea-surface temperature (SST), current speed, mixed-layer depth, and bathymetry. Both zooplankton biomass and the occurrence of fish schools exhibited negative relationships with temperature, and temperature was subsequently an important variable in determining seabird distribution. We were able to establish correlations between the distribution of prey and the spatiotemporal distribution of albatross, little penguins and common-diving petrels. We were unable to find a correlation between the distribution of prey and that of short-tailed shearwaters and fairy prions. For high-use coastal areas, the delineation of important foraging regions is essential to balance human use of an area with the needs of marine predators, particularly seabirds.

Inter- and intrasex habitat partitioning in the highly dimorphic southern elephant seal.

Partitioning resources is a key mechanism for avoiding intraspecific competition and maximizing individual energy gain. However, in sexually dimorphic species it is difficult to discern if partitioning is due to competition or the different resource needs of morphologically distinct individuals. In the highly dimorphic southern elephant seal, there are intersexual differences in habitat use; at Iles Kerguelen, males predominantly use shelf waters, while females use deeper oceanic waters. There are equally marked intrasexual differences, with some males using the nearby Kerguelen Plateau, and others using the much more distant Antarctic continental shelf (~2,000 km away). We used this combination of inter and intrasexual behavior to test two hypotheses regarding habitat partitioning in highly dimorphic species. (a) that intersexual differences in habitat use will not appear until the seals diverge in body size and (b) that some habitats have higher rates of energy return than others. In particular, that the Antarctic shelf would provide higher energy returns than the Kerguelen Shelf, to offset the greater cost of travel. We quantified the habitat use of 187 southern elephant seals (102 adult females and 85 subadult males). The seals in the two groups were the same size (~2.4 m) removing the confounding effect of body size. We found that the intersexual differences in habitat use existed before the divergence in body size. Also, we found that the amount of energy gained was the same in all of the major habitats. This suggests that the use of shelf habitats by males is innate, and a trade-off between the need to access the large benthic prey available on shelf waters, against the higher risk of predation there. Intrasexual differences in habitat use are another trade-off; although there are fewer predators on the Antarctic shelf, it is subject to considerable interannual fluctuations in sea-ice extent. In contrast, the Kerguelen Plateau presents more consistent foraging opportunities, but contains higher levels of predation. Habitat partitioning in this highly dimorphic species is therefore the result of complex interplay of life history strategies, environmental conditions and predation pressure.