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Abstract Background Determining the relative contribution of intrinsic and extrinsic factors to fluctuations in population size, trends and demographic composition is analytically complex. It is often only possible to examine the combined effects of these factors through measurements made over long periods, spanning an array of population densities or levels of food availability. Using age-structured mark-recapture models and datasets spanning five decades (1950–1999), and two periods of differing relative population density, we estimated age-specific probabilities of survival and examined the combined effects of population density and environmental conditions on juvenile survival of southern elephant seals at Macquarie Island. Results First-year survival decreased with density during the period of highest population size, and survival increased during years when the Southern Oscillation Index (SOI) anomaly (deviation from a 50-year mean) during the mother's previous foraging trip to sea was positive (i.e., El Niño). However, when environmental stochasticity and density were considered together, the effect of density on first-year survival effectively disappeared. Ignoring density effects also leads to models placing too much emphasis on the environmental conditions prevailing during the naïve pup's first year at sea. Conclusion Our analyses revealed that both the state of the environment and population density combine to modify juvenile survival, but that the degree to which these processes contributed to the variation observed was interactive and complex. This underlines the importance of evaluating the relative contribution of both the intrinsic and extrinsic factors that regulate animal populations because false conclusions regarding the importance of population regulation may be reached if they are examined in isolation.

Abstract Background Chemical immobilization of Weddell seals (Leptonychotes weddellii) has previously been, for the most part, problematic and this has been mainly attributed to the type of immobilizing agent used. In addition to individual sensitivity, physiological status may play an important role. We investigated the use of the intravenous administration of a 1:1 mixture of tiletamine and zolazepam (Telazol®) to immobilize adult females at different points during a physiologically demanding 5–6 week lactation period. We also compared performance between IV and IM injection of the same mixture. Results The tiletamine:zolazepam mixture administered intravenously was an effective method for immobilization with no fatalities or pronounced apnoeas in 106 procedures; however, there was a 25 % (one animal in four) mortality rate with intramuscular administration. Induction time was slightly longer for females at the end of lactation (54.9 ± 2.3 seconds) than at post-parturition (48.2 ± 2.9 seconds). In addition, the number of previous captures had a positive effect on induction time. There was no evidence for effects due to age, condition (total body lipid), stage of lactation or number of captures on recovery time. Conclusion We suggest that intravenous administration of tiletamine and zolazepam is an effective and safe immobilizing agent for female Weddell seals. Although individual traits could not explain variation in recovery time, we suggest careful monitoring of recovery times during longitudinal studies (> 2 captures). We show that physiological pressures do not substantially affect response to chemical immobilization with this mixture; however, consideration must be taken for differences that may exist for immobilization of adult males and juveniles. Nevertheless, we recommend a mass-specific dose of 0.50 – 0.65 mg/kg for future procedures with adult female Weddell seals and a starting dose of 0.50 mg/kg for other age classes and other phocid seals.

Determining the factors influencing habitat selection and hunting success in top predators is crucial for understanding how these species may respond to environmental changes. For marine top predators, such factors have been documented in pelagic foragers, with habitat use and hunting success being linked to chlorophyll-a concentrations, sea surface temperature and light conditions. In contrast, little is known about the determinants of benthic marine predators. The Australian fur seal (Arctocephalus pusillus doriferus) is a benthic-diving forager that has a breeding and foraging distribution largely restricted to Bass Strait, the shallow (max. depth 80 m) continental shelf region between the Australian mainland and Tasmania. The species forages mostly on benthic prey and represents the greatest resident marine predator biomass in south-eastern Australia. The region is also one of the world's fastest-warming marine areas and oceanographic changes are influencing shifts in prey distribution and abundance. In the present study, GPS-derived locations of benthic dives (n = 288,449) and dive behaviour metrics were used to determine seafloor habitat selection and factors influencing hunting success in 113 lactating adult females from Kanowna Island during the winters of 2006-2021. Individuals non-randomly selected foraging habitats comprised of deeper, steeper sloped, muddy-sandy areas with less gravel and highly disturbed regions (P < 0.01). Hunting success was greatest in shallower rocky reefs (< 30 m) and deep areas (> 40 m) characterised by moderate presence of gravel (25-50%) and substantial rock composition (50-75%) on the seabed. These findings suggest that habitat use and hunting success in adult female Australian fur seals could be impacted by predicted oceanographic changes, such as rising temperature, altered currents and waves which may modify seafloor characteristics and benthic communities., (© 2024. The Author(s).)

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., Competing Interests: There are no competing interests in connection with this manuscript., (© 2023 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., Competing Interests: Conflict of interestThe authors declare that they have no conflict of interest. Author Yongjun Tian is a member of the Editorial Board, but was not involved in the journal’s review of or decision related to this manuscript., (© The Author(s) 2023, corrected publication 2023.)

Plastic pollution is distributed patchily around the world's oceans. Likewise, marine organisms that are vulnerable to plastic ingestion or entanglement have uneven distributions. Understanding where wildlife encounters plastic is crucial for targeting research and mitigation. Oceanic seabirds, particularly petrels, frequently ingest plastic, are highly threatened, and cover vast distances during foraging and migration. However, the spatial overlap between petrels and plastics is poorly understood. Here we combine marine plastic density estimates with individual movement data for 7137 birds of 77 petrel species to estimate relative exposure risk. We identify high exposure risk areas in the Mediterranean and Black seas, and the northeast Pacific, northwest Pacific, South Atlantic and southwest Indian oceans. Plastic exposure risk varies greatly among species and populations, and between breeding and non-breeding seasons. Exposure risk is disproportionately high for Threatened species. Outside the Mediterranean and Black seas, exposure risk is highest in the high seas and Exclusive Economic Zones (EEZs) of the USA, Japan, and the UK. Birds generally had higher plastic exposure risk outside the EEZ of the country where they breed. We identify conservation and research priorities, and highlight that international collaboration is key to addressing the impacts of marine plastic on wide-ranging species., (© 2023. The Author(s).)

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).

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., Competing Interests: We declare no competing interests., (© 2023 The Authors.)

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.

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., (© 2022 The Authors. Global Change Biology published by John Wiley & Sons Ltd.)

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., Competing Interests: The authors have declared that no competing interests exist. The commercial affiliations of authors KP, PGB and KWM does not alter our adherence to PLOS ONE policies on sharing data and materials.

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., Competing Interests: None declared., (© 2022 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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.

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., (© 2021 John Wiley & Sons Ltd.)

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., Competing Interests: The authors declare there are no competing interests., (©2021 Meyers et al.)

The global lockdown to mitigate COVID-19 pandemic health risks has altered human interactions with nature. Here, we report immediate impacts of changes in human activities on wildlife and environmental threats during the early lockdown months of 2020, based on 877 qualitative reports and 332 quantitative assessments from 89 different studies. Hundreds of reports of unusual species observations from around the world suggest that animals quickly responded to the reductions in human presence. However, negative effects of lockdown on conservation also emerged, as confinement resulted in some park officials being unable to perform conservation, restoration and enforcement tasks, resulting in local increases in illegal activities such as hunting. Overall, there is a complex mixture of positive and negative effects of the pandemic lockdown on nature, all of which have the potential to lead to cascading responses which in turn impact wildlife and nature conservation. While the net effect of the lockdown will need to be assessed over years as data becomes available and persistent effects emerge, immediate responses were detected across the world. Thus, initial qualitative and quantitative data arising from this serendipitous global quasi-experimental perturbation highlights the dual role that humans play in threatening and protecting species and ecosystems. Pathways to favorably tilt this delicate balance include reducing impacts and increasing conservation effectiveness., Competing Interests: Authors declare no competing interests., (© 2021 Published by Elsevier Ltd.)

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., (© 2021 The Authors.)

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., Competing Interests: The authors declare there are no competing interests., (©2021 Li et al.)

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., Competing Interests: None declared., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2021 by the Ecological Society of America.)

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., Competing Interests: The authors declare no competing or financial interests., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

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., (© 2021 The Authors. Ecology and Evolution published by John Wiley & Sons Ltd.)

Naturally occurring metals and metalloids [metal(loid)s] are essential for the physiological functioning of wildlife; however, environmental contamination by metal(loid) and plastic pollutants is a health hazard. Metal(loid)s may interact with plastic in the environment and there is mixed evidence about whether plastic ingested by wildlife affects metal(loid) absorption/assimilation and concentration in the body. We examined ingested plastic and liver concentration of eleven metal(loid)s in two seabird species: fairy (Pachyptila turtur) and slender-billed prions (P. belcheri). We found significant relationships between ingested plastic and the concentrations of aluminium (Al), manganese (Mn), iron (Fe), cobalt (Co), copper (Cu) and zinc (Zn) in the liver of prions. We investigated whether the pattern of significant relationships reflected plastic-metal(loid) associations predicted in the scientific literature, including by transfer of metals from ingested plastics or malnutrition due to dietary dilution by plastics in the gut. We found some support for both associations, suggesting that ingested plastic may be connected with dietary dilution / lack of essential nutrients, especially iron, and potential transfer of zinc. We did not find a relationship between plastic and non-essential metal(loid)s, including lead. The effect of plastic was minor compared to that of dietary exposure to metal(oid)s, and small plastic loads (< 3 items) had no discernible link with metal(loid)s. This new evidence shows a relationship between plastic ingestion and liver metal(loid) concentrations in free-living wildlife.

Understanding the factors which influence foraging behaviour and success in marine mammals is crucial to predicting how their populations may respond to environmental change. The Australian fur seal (Arctocephalus pusillus doriferus, AUFS) is a predominantly benthic forager on the shallow continental shelf of Bass Strait, and represents the greatest biomass of marine predators in south-eastern Australia. The south-east Australian region is experiencing rapid oceanic warming, predicted to lead to substantial alterations in prey diversity, distribution and abundance. In the present study, foraging effort and indices of foraging success and efficiency were investigated in 138 adult female AUFS (970 foraging trips) during the winters of 1998-2019. Large scale climate conditions had a strong influence on foraging effort, foraging success and efficiency. Foraging effort and foraging success were also strongly influenced by winter chlorophyll-a concentrations and sea-surface height anomalies in Bass Strait. The results suggest increasing foraging effort and decreasing foraging success and efficiency under anticipated environmental conditions, which may have population-level impacts.

Changes in the foraging environment and at-sea distribution of southern elephant seals from Kerguelen Islands were investigated over a decade (2004-2018) using tracking, weaning mass, and blood δ 13 C values. Females showed either a sub-Antarctic or an Antarctic foraging strategy, and no significant shift in their at-sea distribution was detected between 2004 and 2017. The proportion of females foraging in sub-Antarctic versus Antarctic habitats did not change over the 2006-2018 period. Pup weaning mass varied according to the foraging habitat of their mothers. The weaning mass of sub-Antarctic foraging mothers' pups decreased by 11.7 kg over the study period, but they were on average 5.8 kg heavier than pups from Antarctic foraging mothers. Pup blood δ 13 C values decreased by 1.1‰ over the study period regardless of their sex and the presumed foraging habitat of their mothers. Together, these results suggest an ecological change is occurring within the Indian sector of the Southern Ocean with possible consequences on the foraging performance of southern elephant seals. We hypothesize that this shift in δ 13 C is related to a change in primary production and/or in the composition of phytoplankton communities, but this requires further multidisciplinary investigations.

Background: State-space models are important tools for quality control and analysis of error-prone animal movement data. The near real-time (within 24 h) capability of the Argos satellite system can aid dynamic ocean management of human activities by informing when animals enter wind farms, shipping lanes, and other intensive use zones. This capability also facilitates the use of ocean observations from animal-borne sensors in operational ocean forecasting models. Such near real-time data provision requires rapid, reliable quality control to deal with error-prone Argos locations., Methods: We formulate a continuous-time state-space model to filter the three types of Argos location data (Least-Squares, Kalman filter, and Kalman smoother), accounting for irregular timing of observations. Our model is deliberately simple to ensure speed and reliability for automated, near real-time quality control of Argos location data. We validate the model by fitting to Argos locations collected from 61 individuals across 7 marine vertebrates and compare model-estimated locations to contemporaneous GPS locations. We then test assumptions that Argos Kalman filter/smoother error ellipses are unbiased, and that Argos Kalman smoother location accuracy cannot be improved by subsequent state-space modelling., Results: Estimation accuracy varied among species with Root Mean Squared Errors usually <5 km and these decreased with increasing data sampling rate and precision of Argos locations. Including a model parameter to inflate Argos error ellipse sizes in the north - south direction resulted in more accurate location estimates. Finally, in some cases the model appreciably improved the accuracy of the Argos Kalman smoother locations, which should not be possible if the smoother is using all available information., Conclusions: Our model provides quality-controlled locations from Argos Least-Squares or Kalman filter data with accuracy similar to or marginally better than Argos Kalman smoother data that are only available via fee-based reprocessing. Simplicity and ease of use make the model suitable both for automated quality control of near real-time Argos data and for manual use by researchers working with historical Argos data., Competing Interests: Competing interestsThe authors declare that they have no competing interests., (© The Author(s) 2020.)

Invasive species present a major conservation threat globally and nowhere are their affects more pronounced than in island ecosystems. Determining how native island populations respond demographically to invasive species can provide information to mitigate the negative effects of invasive species. Using 20 years of mark-recapture data from three sympatric species of albatrosses (black-browed Thalassarche melanophris, grey-headed T. chrysostoma, and light-mantled albatrosses Phoebetria palpebrata), we quantified the influence of invasive European rabbits Oryctolagus cuniculus and extreme weather patterns on breeding probability and success. Temporal variability in rabbit density explained 33-76% of the variability in breeding probability for all three species, with severe decreases in breeding probability observed after a lag period following highest rabbit numbers. For black-browed albatrosses, the combination of extreme rainfall and high rabbit density explained 33% of total trait variability and dramatically reduced breeding success. We showed that invasive rabbits and extreme weather events reduce reproductive output in albatrosses and that eliminating rabbits had a positive effect on albatross reproduction. This illustrates how active animal management at a local breeding site can result in positive population outcomes even for wide ranging animals like albatrosses where influencing vital rates during their at-sea migrations is more challenging.

Southern Ocean ecosystems are under pressure from resource exploitation and climate change 1,2 . 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 Retrospective Analysis of Antarctic Tracking Data (RAATD) is a Scientific Committee for Antarctic Research project led jointly by the Expert Groups on Birds and Marine Mammals and Antarctic Biodiversity Informatics, and endorsed by the Commission for the Conservation of Antarctic Marine Living Resources. RAATD consolidated tracking data for multiple species of Antarctic meso- and top-predators to identify Areas of Ecological Significance. These datasets and accompanying syntheses provide a greater understanding of fundamental ecosystem processes in the Southern Ocean, support modelling of predator distributions under future climate scenarios and create inputs that can be incorporated into decision making processes by management authorities. In this data paper, we present the compiled tracking data from research groups that have worked in the Antarctic since the 1990s. The data are publicly available through biodiversity.aq and the Ocean Biogeographic Information System. The archive includes tracking data from over 70 contributors across 12 national Antarctic programs, and includes data from 17 predator species, 4060 individual animals, and over 2.9 million observed locations.

Mesopelagic fish and squid occupy ocean depths extending below the photic zone and their vertical migrations represent a massive pathway moving energy and carbon through the water column. Their spatio-temporal distribution is however, difficult to map across remote regions particularly the vast Southern Ocean. This represents a key gap in understanding biogeochemical processes, marine ecosystem structure, and how changing ocean conditions will affect marine predators, which depend upon mesopelagic prey. We infer mesopelagic prey vertical distribution and relative abundance in the Indian sector of the Southern Ocean (20° to 130°E) with a novel approach using predator-derived indices. Fourteen years of southern elephant seal tracking and dive data, from the open ocean between the Antarctic Polar Front and the southern Antarctic Circumpolar Current front, clearly show that the vertical distribution of mesopelagic prey is influenced by the physical hydrographic processes that structure their habitat. Mesopelagic prey have a more restricted vertical migration and higher relative abundance closer to the surface where Circumpolar Deep Water rises to shallower depths. Combining these observations with a future projection of Southern Ocean conditions we show that changes in the coupling of surface and deep waters will potentially redistribute mesopelagic prey. These changes are small overall, but show important spatial variability: prey will increase in relative abundance to the east of the Kerguelen Plateau but decrease to the west. The consequences for deep-diving specialists such as elephant seals and whales over this time scale will likely be minor, but the changes in mesoscale vertical energy flow have implications for predators that forage within the mesopelagic zone as well as the broader pelagic ecosystem.

Background: To understand and predict the distribution of foragers, it is crucial to identify the factors that affect individual movement decisions at different scales. Individuals are expected to adjust their foraging movements to the hierarchical spatial distribution of resources. At a small local scale, spatial segregation in foraging habitat happens among individuals of closely situated colonies. If foraging segregation is due to differences in distribution of resources, we would expect segregated foraging areas to have divergent habitat characteristics., Results: We investigated how environmental characteristics of preferred foraging areas differ between two closely situated Subantarctic fur seal (Arctocephalus tropicalis) colonies and a single Antarctic fur seal (A. gazella) colony that forage in different pelagic areas even though they are located well within each other's foraging range. We further investigated the influence of the seasonal cycle on those environmental factors. This study used tracking data from 121 adult female Subantarctic and Antarctic fur seals, collected during summer and winter (2009-2015), from three different colonies. Boosted Regression Tree species distribution models were used to determine key environmental variables associated with areas of fur seal restricted search behaviour. There were no differences in the relative influence of key environmental variables between colonies and seasons. The variables with the most influence for each colony and season were latitude, longitude and magnitude of sea-currents. The influence of latitude and longitude is a by-product of the species' distinct foraging areas, despite the close proximity (< 25 km) of the colonies. The predicted potential foraging areas for each colony changed from summer to winter, reflecting the seasonal cycle of the Southern Ocean. The model predicted that the potential foraging areas of females from the three colonies should overlap, and the fact they do not in reality indicates that factors other than environmental are influencing the location of each colony's foraging area., Conclusions: The results indicated that small scale spatial segregation of foraging habitats is not driven by bottom-up processes. It is therefore important to also consider other potential drivers, e.g. competition, information transfer, and memory, to understand animal foraging decisions and movements.

Foraging behaviour of marine predators inferred from the analysis of horizontal or vertical movements commonly lack quantitative information about foraging success. Several marine mammal species are known to perform dives where they passively drift in the water column, termed "drift" dives. The drift rate is determined by the animal's buoyancy, which can be used to make inference regarding body condition. Long term dive records retrieved via satellite uplink are often summarized before transmission. This loss of resolution hampers identification of drift dives. Here, we develop a flexible, hierarchically structured approach to identify drift dives and estimate the drift rate from the summarized time-depth profiles that are increasingly available to the global research community. Based on high-resolution dive data from southern elephant seals, we classify dives as drift/non-drift and apply a summarization algorithm. We then (i) automatically generate dive groups based on inflection point ordering using a 'Reverse' Broken-Stick Algorithm, (ii) develop a set of threshold criteria to apply across groups, ensuring non-drift dives are most efficiently rejected, and (iii) finally implement a custom Kalman filter to retain the remaining dives that are within the seals estimated drifting time series. Validation with independent data sets shows our method retains approximately 3% of all dives, of which 88% are true drift dives. The drift rate estimates are unbiased, with the upper 95% quantile of the mean squared error between the daily averaged summarized profiles using our method (SDDR) and the observed daily averaged drift rate (ODDR) being only 0.0015. The trend of the drifting time-series match expectations for capital breeders, showing the lowest body condition commencing foraging trips and a progressive improvement as they remain at sea. Our method offers sufficient resolution to track small changes in body condition at a fine temporal scale. This approach overcomes a long-term challenge for large existing and ongoing data collections, with potential application across other drift diving species. Enabling robust identification of foraging success at sea offers a rare and valuable opportunity for monitoring marine ecosystem productivity in space and time by tracking the success of a top predator.

Pollution of the world's oceans by marine debris has direct consequences for wildlife, with fragments of plastic <10 mm the most abundant buoyant litter in the ocean. Seabirds are susceptible to debris ingestion, commonly mistaking floating plastics for food. Studies have shown that half of petrel species regularly ingest anthropogenic waste. Despite the regularity of debris ingestion, no studies to date have quantified the dimensions of debris items ingested across petrel species ranging in size. We excised and measured 1694 rigid anthropogenic debris items from 348 petrel carcasses of 20 species. We found that although the size of items ingested by petrels scale positively with the size of the bird, 90% of all debris items ingested across species fall within a narrow "danger zone" range of 2-10 mm, overlapping with the most abundant oceanic debris size. We conclude that this globally profuse size range of marine plastics is an ingestion hazard to petrels., (Crown Copyright © 2019. Published by Elsevier Ltd. All rights reserved.)

Age structure is a fundamental aspect of animal population biology. Age is strongly related to individual physiological condition, reproductive potential and mortality rate. Currently, there are no robust molecular methods for age estimation in birds. Instead, individuals must be ringed as chicks to establish known-age populations, which is a labour-intensive and expensive process. The estimation of chronological age using DNA methylation (DNAm) is emerging as a robust approach in mammals including humans, mice and some non-model species. Here, we quantified DNAm in whole blood samples from a total of 71 known-age Short-tailed shearwaters (Ardenna tenuirostris) using digital restriction enzyme analysis of methylation (DREAM). The DREAM method measures DNAm levels at thousands of CpG dinucleotides throughout the genome. We identified seven CpG sites with DNAm levels that correlated with age. A model based on these relationships estimated age with a mean difference of 2.8 years to known age, based on validation estimates from models created by repeated sampling of training and validation data subsets. Longitudinal observation of individuals re-sampled over 1 or 2 years generally showed an increase in estimated age (6/7 cases). For the first time, we have shown that epigenetic changes with age can be detected in a wild bird. This approach should be of broad interest to researchers studying age biomarkers in non-model species and will allow identification of markers that can be assessed using targeted techniques for accurate age estimation in large population studies., (© 2018 John Wiley & Sons Ltd.)

Procellariiformes are the most threatened bird group globally, and the group with the highest frequency of marine debris ingestion. Marine debris ingestion is a globally recognized threat to marine biodiversity, yet the relationship between how much debris a bird ingests and mortality remains poorly understood. Using cause of death data from 1733 seabirds of 51 species, we demonstrate a significant relationship between ingested debris and a debris-ingestion cause of death (dose-response). There is a 20.4% chance of lifetime mortality from ingesting a single debris item, rising to 100% after consuming 93 items. Obstruction of the gastro-intestinal tract is the leading cause of death. Overall, balloons are the highest-risk debris item; 32 times more likely to result in death than ingesting hard plastic. These findings have significant implications for quantifying seabird mortality due to debris ingestion, and provide identifiable policy targets aimed to reduce mortality for threatened species worldwide.

The emergence of longline fishing around the world has been concomitant with an increase in depredation-interactions by odontocete whales (removal of fish caught on hooks), resulting in substantial socio-economic and ecological impacts. The extent, trends and underlying mechanisms driving these interactions remain poorly known. Using long-term (2003-2017) datasets from seven major Patagonian toothfish (Dissostichus eleginoides) longline fisheries, this study assessed the levels and inter-annual trends of sperm whale (Physeter macrocephalus) and/or killer whale (Orcinus orca) interactions as proportions of fishing time (days) and fishing area (spatial cells). The role of fishing patterns in explaining between-fisheries variations of probabilities of odontocete interactions was investigated. While interaction levels remained globally stable since the early 2000s, they varied greatly between fisheries from 0 to >50% of the fishing days and area. Interaction probabilities were influenced by the seasonal concentration of fishing effort, size of fishing areas, density of vessels, their mobility and the depth at which they operated. The results suggest that between-fisheries variations of interaction probabilities are largely explained by the extent to which vessels provide whales with opportunities for interactions. Determining the natural distribution of whales will, therefore, allow fishers to implement better strategies of spatio-temporal avoidance of depredation.

Like many species, movement patterns of southern elephant seals (Mirounga leonina) are being influenced by long-term environmental change. These seals migrate up to 4,000 km from their breeding colonies, foraging for months in a variety of Southern Ocean habitats. Understanding how movement patterns vary with environmental features and how these relationships differ among individuals employing different foraging strategies can provide insight into foraging performance at a population level. We apply new fast-estimation tools to fit mixed effects within a random walk movement model, rapidly inferring among-individual variability in southern elephant seal environment-movement relationships. We found that seals making foraging trips to the sea ice on or near the Antarctic continental shelf consistently reduced speed and directionality (move persistence) with increasing sea-ice coverage but had variable responses to chlorophyll a concentration, whereas seals foraging in the open ocean reduced move persistence in regions where circumpolar deep water shoaled. Given future climate scenarios, open-ocean foragers may encounter more productive habitat but sea-ice foragers may see reduced habitat availability. Our approach is scalable to large telemetry data sets and allows flexible combinations of mixed effects to be evaluated via model selection, thereby illuminating the ecological context of animal movements that underlie habitat usage., (© 2018 by the Ecological Society of America.)

Over the past five decades, marine mammal interactions with fisheries have become a major human-wildlife conflict globally. The emergence of longline fishing is concomitant with the development of depredation-type interactions i.e., marine mammals feeding on fish caught on hooks. The killer whale ( Orcinus orca ) is one of the species most involved in depredation on longline fisheries. The issue was first reported in high latitudes but, with increasing expansion of this fishing method, other fisheries have begun to experience interactions. The present study investigated killer whale interactions with two geographically isolated blue-eye trevalla ( Hyperoglyphe antarctica ) fisheries operating in temperate waters off Amsterdam/St. Paul Islands (Indian Ocean) and south-eastern Australia. These two fisheries differ in the fishing technique used (vertical vs. demersal longlines), effort, catch, fleet size and fishing area size. Using 7-year (2010-16) long fishing and observation datasets, this study estimated the levels of killer whale interactions and examined the influence of spatio-temporal and operational variables on the probability of vessels to experience interactions. Killer whales interactions occurred during 58.4% and 21.2% of all fishing days, and over 94% and 47.4% of the fishing area for both fisheries, respectively. In south-eastern Australia, the probability of occurrence of killer whale interactions during fishing days varied seasonally with a decrease in spring, increased with the daily fishing effort and decreased with the distance travelled by the vessel between fishing days. In Amsterdam/St. Paul, this probability was only influenced by latitude, with an increase in the southern part of the area. Together, these findings document two previously unreported cases of high killer whale depredation, and provide insights on ways to avoid the issue. The study also emphasizes the need to further examine the local characteristics of fisheries and the ecology of local depredating killer whale populations in as important drivers of depredation., Competing Interests: The authors declare there are no competing interests.

Higher trophic-level species are an integral component of any marine ecosystem. Despite their importance, methods for representing these species in end-to-end ecosystem models often have limited representation of life histories, energetics and behaviour. We built an individual-based model coupled with a dynamic energy budget for female southern elephant seals Mirounga leonina to demonstrate a method for detailed representation of marine mammals. We aimed to develop a model which could i) simulate energy use and life histories, as well as breeding traits of southern elephant seals in an emergent manner, ii) project a stable population over time, and iii) have realistic population dynamics and structure based on emergent life history features (such as age at first breeding, lifespan, fecundity and (yearling) survival). We evaluated the model's ability to represent a stable population over long time periods (>10 generations), including the sensitivity of the emergent properties to variations in key parameters. Analyses indicated that the model is sensitive to changes in resource availability and energy requirements for the transition from pup to juvenile, and juvenile to adult stage. This was particularly the case for breeding success and yearling survival. This model is suitable for use as a standalone tool for investigating the impacts of changes to behaviour and population responses of southern elephant seals.

Antarctic coastal polynyas are regions of persistent open water and are thought to be key bio-physical features within the sea-ice zone. However, their use by the upper trophic levels of ecosystems remains unclear. A unique bio-physical dataset recorded by southern elephant seals reveals that East Antarctic polynyas are a key winter foraging habitat for male seals. During their post-moult trips from Isles Kerguelen to the Antarctic continental shelf, a total of 18 out of 23 seals visited 9 different polynyas, spending on average 25 ± 20% (up to 75%) of their total trip time inside polynyas. Changes in seal foraging and diving behaviours are observed inside polynyas as compared to outside polynyas. Two polynya usages by seals are observed for the inactive and active polynya phases, pointing to different seasonal peaks in prey abundance. During the active polynya phase, we link seal foraging behaviour to changes in the physical stability of the water-column, which likely impact the seasonal biological dynamics within polynyas.

In iteroparous species, intermittent breeding is an important life-history tactic that can greatly affect animal population growth and viability. Despite its importance, few studies have quantified the consequences of breeding pauses on lifetime reproductive output, principally because calculating lifetime reproductive output requires knowledge of each individual's entire reproductive history. This information is extremely difficult to obtain in wild populations. We applied novel statistical approaches that account for uncertainty in state assessment and individual heterogeneity to an 18-year capture-recapture dataset of 6,631 female southern elephant seals from Macquarie Island. We estimated survival and breeding probabilities, and investigated the consequences of intermittent breeding on lifetime reproductive output. We found consistent differences in females' demographic performance between two heterogeneity classes. In particular, breeding imbued a high cost on survival in the females from the heterogeneity class 2, assumed to be females of lower quality. Individual quality also appeared to play a major role in a female's decision to skip reproduction with females of poorer quality more likely to skip breeding events than females of higher quality. Skipping some breeding events allowed females from both heterogeneity classes to increase lifetime reproductive output over females that bred annually. However, females of lower quality produced less offspring over their lifetime. Intermittent breeding seems to be used by female southern elephant seals as a tactic to offset reproductive costs on survival and enhance lifetime reproductive output but remains unavoidable and driven by individual-specific constraints in some other females., (© 2017 The Authors. Journal of Animal Ecology © 2017 British Ecological Society.)

Most seabirds do not have any outward identifiers of their chronological age, so estimation of seabird population age structure generally requires expensive, long-term banding studies. We investigated the potential to use a molecular age biomarker to estimate age in short-tailed shearwaters (Ardenna tenuirostris). We quantified DNA methylation in several A. tenuirostris genes that have shown age-related methylation changes in mammals. In birds ranging from chicks to 21 years of age, bisulphite treated blood and feather DNA was sequenced and methylation levels analysed in 67 CpG sites in 13 target gene regions. From blood samples, five of the top relationships with age were identified in KCNC3 loci (CpG66: R2 = 0.325, p = 0.019). In feather samples ELOVL2 (CpG42: R2 = 0.285, p = 0.00048) and EDARADD (CpG46: R2 = 0.168, p = 0.0067) were also weakly correlated with age. However, the majority of markers had no clear association with age (of 131 comparisons only 12 had a p-value < 0.05) and statistical analysis using a penalised lasso approach did not produce an accurate ageing model. Our data indicate that some age-related signatures identified in orthologous mammalian genes are not conserved in the long-lived short tailed shearwater. Alternative molecular approaches will be required to identify a reliable biomarker of chronological age in these seabirds.

Understanding divergent biological responses to climate change is important for predicting ecosystem level consequences. We use species habitat models to predict the winter foraging habitats of female southern elephant seals and investigate how changes in environmental variables within these habitats may be related to observed decreases in the Macquarie Island population. There were three main groups of seals that specialized in different ocean realms (the sub-Antarctic, the Ross Sea and the Victoria Land Coast). The physical and climate attributes (e.g. wind strength, sea surface height, ocean current strength) varied amongst the realms and also displayed different temporal trends over the last two to four decades. Most notably, sea ice extent increased on average in the Victoria Land realm while it decreased overall in the Ross Sea realm. Using a species distribution model relating mean residence times (time spent in each 50 × 50 km grid cell) to 9 climate and physical co-variates, we developed spatial predictions of residence time to identify the core regions used by the seals across the Southern Ocean from 120°E to 120°W. Population size at Macquarie Island was negatively correlated with ice concentration within the core habitat of seals using the Victoria Land Coast and the Ross Sea. Sea ice extent and concentration is predicted to continue to change in the Southern Ocean, having unknown consequences for the biota of the region. The proportion of Macquarie Island females (40%) utilizing the relatively stable sub-Antarctic region, may buffer this population against longer-term regional changes in habitat quality, but the Macquarie Island population has persistently decreased (-1.45% per annum) over seven decades indicating that environmental changes in the Antarctic are acting on the remaining 60% of the population to impose a long-term population decline in a top Southern Ocean predator., (© 2017 John Wiley & Sons Ltd.)

DNA methylation (DNAm) is a key mechanism for regulating gene expression in animals and levels are known to change with age. Recent studies have used DNAm changes as a biomarker to estimate chronological age in humans and these techniques are now also being applied to domestic and wild animals. Animal age is widely used to track ongoing changes in ecosystems, however chronological age information is often unavailable for wild animals. An ability to estimate age would lead to improved monitoring of (i) population trends and status and (ii) demographic properties such as age structure and reproductive performance. Recent studies have revealed new examples of DNAm age association in several new species increasing the potential for developing DNAm age biomarkers for a broad range of wild animals. Emerging technologies for measuring DNAm will also enhance our ability to study age-related DNAm changes and to develop new molecular age biomarkers.

Predicting population declines is a key challenge in the face of global environmental change. Abundance-based early warning signals have been shown to precede population collapses; however, such signals are sensitive to the low reliability of abundance estimates. Here, using historical data on whales harvested during the 20th century, we demonstrate that early warning signals can be present not only in the abundance data, but also in the more reliable body size data of wild populations. We show that during the period of commercial whaling, the mean body size of caught whales declined dramatically (by up to 4 m over a 70-year period), leading to early warning signals being detectable up to 40 years before the global collapse of whale stocks. Combining abundance and body size data can reduce the length of the time series required to predict collapse, and decrease the chances of false positive early warning signals.

The population of Southern elephant seals (Mirounga leonina) at Macquarie Island has declined since the 1960s, and is thought to be due to changing oceanic conditions leading to reductions in the foraging success of Macquarie Island breeding females. To test this hypothesis, we used a 55-year-old data set on weaning size of southern elephant seals to quantify a decrease in weaning size from a period of population stability in 1950s to its present state of on-going decline. Being capital breeders, the size of elephant seal pups at weaning is a direct consequence of maternal foraging success in the preceding year. During the 1940-1950s, the mean of female pups at weaning was similar between the Heard and Macquarie Island populations, while the snout-tail-length length of male weaners from Heard Island were longer than their conspecifics at Macquarie Island. Additionally, the snout-tail-length of pups at weaning decreased by 3cm between the 1950s and 1990s in the Macquarie Island population, concurrent with the observed population decline. Given the importance of weaning size in determining first-year survival and recruitment rates, the decline in the size at weaning suggests that the decline in the Macquarie Island population has, to some extent, been driven by reduced maternal foraging success, consequent declines in the size of pups at weaning, leading to reduced first-year survival rates and recruitment of breeding females into the population 3 to 4 years later.

Cetacean energy stores are known to vary according to life history, reproductive status and time of year; however, the opportunity to quantify these relationships is rare. Using a unique set of historical whaling records from Western Australia (1952-1963), we investigated energy stores of large cetaceans with differing life histories, and quantified the relationship between total body lipid and length for humpback whales ( Megaptera novaeangliae) ( n  = 905) and sperm whales (Physeter macrocephalus) ( n  = 1961). We found that total body lipid increased with body length in both humpback and sperm whales, consistent with size-related energy stores. Male humpback whales stored 2.49 kl (15.6 barrels) (31.9-74.9%) more lipid than male sperm whales of equivalent length, to fuel their annual migration. Relative lipid stores of sperm whales (males) were constant throughout the year, while those of humpback whales varied with reproductive class and sampling date. Pregnant female humpback whales had higher relative energy stores than non-pregnant females and males (26.2% and 37.4%, respectively), to fuel the energy demands of gestation and lactation. Those that reached the sampling site later ( en route to their breeding grounds) carried higher lipid stores than those that arrived earlier, possibly reflecting individual variation in residency times in the Antarctic feeding grounds. Importantly, longer pregnant females had relatively larger energy stores than the shorter pregnant females, indicating that the smaller individuals may experience higher levels of energetic stress during the migration fast. The relationships we developed between body lipid and length can be used to inform bioenergetics and ecosystem models when such detailed information is not available.