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Understanding how closely related, sympatric species distribute themselves relative to their environment is critical to understanding ecosystem structure and function and predicting effects of environmental variation. The Antarctic Peninsula supports high densities of krill and krill consumers; however, the region is warming rapidly, with unknown consequences. Humpback whales Megaptera novaeangliae and Antarctic minke whales Balaenoptera bonaerensis are the largest krill consumers here, yet key data gaps remain about their distribution, behavior, and interactions and how these will be impacted by changing conditions. Using satellite telemetry and novel spatial point-process modeling techniques, we quantified habitat use of each species relative to dynamic environmental variables and determined overlap in core habitat areas during summer months when sea ice is at a minimum. We found that humpback whales ranged broadly over continental shelf waters, utilizing nearshore bays, while minke whales restricted their movements to sheltered bays and areas where ice is present. This presents a scenario where minke whale core habitat overlaps substantially with the broader home ranges of humpback whales. While there is no indication that prey is limiting in this ecosystem, increased overlap between these species may arise as climate-driven changes that affect the extent, timing, and duration of seasonal sea ice decrease the amount of preferred foraging habitat for minke whales while concurrently increasing it for humpback whales. Our results provide the first quantitative assessment of behaviorally based habitat use and sympatry between these 2 krill consumers and offers insight into the potential effects of a rapidly changing environment on the structure and function of a polar ecosystem.

There is potential value in exploring multi-stock models to address situations where humpback stocks are mixing. However, sensitivity to the assumptions underlying these models has yet to be fully explored. Using a simple simulation approach, the assumptions of a population model that allows for mixing of humpback whale (Megaptera novaeangliae) stocks D and E on feeding areas has been explored by relaxing the assumptions of the original Johnston and Butterworth model in a number of plausible ways. First the ability of the model to estimate parameters was checked for a situation where simulated data are generated from an underlying model of exactly the same form for which the actual values of these parameters are known (Scenario 1). Then the ability of the model to estimate these parameters when alternative forms and assumptions were used for the underlying model generating the data was investigated. Specifically, stocks were allowed to mix non-uniformly across each feeding area and catch was non-uniformly distributed across each feeding area (Scenario 2). The consequences of density dependence implemented on feeding rather than breeding areas (Scenario 3) were also examined. The original mixing model was robust to alternate mixing and catch allocation scenarios in all but one of the simulations, but when density dependence acted at the level of the feeding rather than the breeding areas, the model produced estimates that were quite different from the underlying population. It is recommend that the inclusion of density dependence on feeding areas in models that allow for mixing of whales on these grounds be investigated further.

Many agencies provided funding and logistical support for the various research efforts resulting in the data presented here: the South African Department of Science and Technology, administered by the National Research Foundation and the Department of Environmental Affairs for subantarctic fur seal work; the Australian Research Council (DP110102065), Holsworth Wildlife Research Endowment and the Office of Naval Research (Marine Mammals and Biological Oceanography Program Award no. N00014-10-1-0385) for Australian fur seal work; the National Oceanic and Atmospheric Administration (NOAA) via grants to the Alaska SeaLife Center and the National Marine Mammal Laboratory, with additional funding and logistical support from North Pacific Wildlife Consulting for Steller sea lion and northern fur seal (Russia) work; the National Marine Mammal Laboratory, National Marine Fisheries Service, NOAA for northern fur seal (Alaska) work. Research support for R.W. Davis was provided by the National Science Foundation. Management of gases during diving is not well understood across marine mammal species. Prior to diving, phocid (true) seals generally exhale, a behaviour thought to assist with the prevention of decompression sickness. Otariid seals (fur seals and sea lions) have a greater reliance on their lung oxygen stores, and inhale prior to diving. One otariid, the Antarctic fur seal (Arctocephalus gazella), then exhales during the final 50–85% of the return to the surface, which may prevent another gas management issue: shallow-water blackout. Here, we compare data collected from animal-attached tags (video cameras, hydrophones and conductivity sensors) deployed on a suite of otariid seal species to examine the ubiquity of ascent exhalations for this group. We find evidence for ascent exhalations across four fur seal species, but that such exhalations are absent for three sea lion species. Fur seals and sea lions are no longer genetically separated into distinct subfamilies, but are morphologically distinguished by the thick underfur layer of fur seals. Together with their smaller size and energetic dives, we suggest their air-filled fur might underlie the need to perform these exhalations, although whether to reduce buoyancy and ascent speed, for the avoidance of shallow-water blackout or to prevent other cardiovascular management issues in their diving remains unclear. Postprint Postprint

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

The Antarctic krill fishery is the largest in the southern ocean, but currently operates without fine-scale information on whale movement and behavior. Using a multi-year dataset of satellite-tagged whales, as well as information on krill catch levels, we analyzed the spatial distribution of whales and fisheries effort within the small-scale management units defined by the Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR). Using a Bayesian movement model to partition whale movement into traveling and area-restricted search states, we found that both whale behavior and krill catch effort were spatially clustered, with distinct hotspots of the whale activity in the Gerlache and southern Branfield Straits. These areas align with increases in krill fishing effort, and present potential areas of current and future conflict. We recommend that the Antarctic West and Bransfield Strait West management units merit particular attention when setting fine-scale catch limits and, more broadly, consideration as critical areas for krill predator foraging.

Satellite telemetry data was used to predict at sea spatial usage of five top order and meso-predators; Antarctic fur seals ( Arctocephalus gazella ), macaroni penguins ( Eudyptes chrysolophus ), king penguins ( Aptenodytes patagonicus ), black browed albatross ( Diomedea melanophrys ), and light mantled albatross ( Phoebetria palpebrata ). All were tagged at Heard Island in the Southern Ocean over a single summer season collecting over 5000 tracking days for 178 individuals. We aimed to predict areas of likely high foraging value from tracking environmental data and also to quantify overlap in foraging range between species. Hidden Markov models were used to differentiate between bouts of Area Restricted Search (ARS) assumed to be associated with areas of higher foraging value, and transit behaviours. Oceanographic and distance metrics associated with ARS activity were then used to calculate a habitat electivity index. A combined bootstrap/Monte Carlo scheme was employed to propagate uncertainty from the Hidden Markov models into the habitat prediction scheme. Distinct differences were predicted in the spatial distribution of foraging locations in different species, reflecting different dispersive abilities and foraging strategy. The extent of usage and foraging distribution was largely contained within Australian the Economic Exclusion Zone (EEZ). In comparison, the smaller Australian Commonwealth Marine Protected Areas (MPAs) contained

Animal movement research relies on biotelemetry, and telemetry‐based locations are increasingly augmented with ancillary information. This presents an underutilized opportunity to enhance movement process models. Given tags designed to record specific behaviors, efforts are increasing to update movement models beyond reliance solely upon horizontal movement information to improve inference of space use and activity budgets. We present two state‐space models adapted to incorporate ancillary data to inform three discrete movement states: directed, resident, and an activity state. These were developed for two case studies: (1) a “haulout” model for Weddell seals, and (2) an “activity” model for Antarctic fur seals which intersperse periods of diving activity and inactivity. The methodology is easily implementable with any ancillary data that can be expressed as a proportion (or binary) indicator. A comparison of the models augmented with ancillary information and unaugmented models confirmed that many behavioral states appeared mischaracterized in the latter. Important changes in subsequent activity budgets occurred. Haulout accounted for 0.17 of the overall Weddell seal time budget, with the estimated proportion of time spent in a resident state reduced from a posterior median of 0.69 (0.65–0.73; 95% HPDI) to 0.54 (0.50–0.58 HPDI). The drop was more dramatic in the Antarctic fur seal case, from 0.57 (0.52–0.63 HPDI) to 0.22 (0.20–0.25 HPDI), with 0.35 (0.31–0.39 HPDI) of time spent in the inactive (nondiving) state. These findings reinforce previously raised contentions about the drawbacks of behavioral states inferred solely from horizontal movements. Our findings have implications for assessing habitat requirements; estimating energetics and consumption; and management efforts such as mitigating fisheries interactions. Combining multiple sources of information within integrated frameworks should improve inference of relationships between movement decisions and fitness, the interplay between resource and habitat dependencies, and their changes at the population and landscape level.

The Antarctic marine environment is changing, and changes in the Southwest Atlantic sector have included decreases in sea ice and increases in water temperature. Associated with these changes is a reported 38% and 81% per decade decline in the numerical density (hereafter density) of Antarctic krill Euphausia superba Dana, 1850 , between 1976 and 2003. Few changes in other components of the ecosystem that could be attributed to such a change, such as a mass decline in krill-dependent predators, have been detected. In an ecosystem so dependent on this keystone species, a massive population decline in krill ought to have had an obvious effect. In the absence of such an effect, it is timely to revisit the issue of the purported decline in krill density. The original analysis that indicated a decline in krill density was based on the 2004 version of KRILLBASE, a database of net samples. We analysed the publicly available and updated version (version 1, accessed 30 November 2017) and our analyses did not suggest a significant decline in krill density. Rather, after accounting for sampling heterogeneity and habitat variables, average krill density appears to have been stable but with considerable inter-annual variability. Since our results were unable to find any evidence for a decline in krill density we recommend a re-appraisal of many of the paradigms that underlie much of the recent thinking about ecosystem change Antarctic waters. Such a revision is necessary to provide a firmer foundation for predictions of the effects of climate change and resource extraction on the Southern Ocean ecosystem.

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., Measurement(s)geographic locationTechnology Type(s)digital curationFactor Type(s)temporal interval • geographic locationSample Characteristic – OrganismAptenodytes forsteri • Lobodon carcinophaga • Eudyptes chrysolophus • Thalassarche melanophrys • Phoebetria palpebrata • Aptenodytes patagonicus • Pygoscelis adeliae • Leptonychotes weddellii • Megaptera novaeangliae • Mirounga leonina • Arctocephalus gazella • Thalassoica antarctica • Eudyptes schlegeli • Diomedea exulans • Phoebetria fusca • Diomedea chrysostoma • Procellaria aequinoctialisSample Characteristic – Environmentpolar biomeSample Characteristic – LocationAntarctica Machine-accessible metadata file describing the reported data: 10.6084/m9.figshare.11688876

Humpback whale (Megaptera novaeangliae) populations typically undertake seasonal migrations, spending winters in low latitude breeding grounds and summers foraging in high latitude feeding grounds. Until recently, a broad scale understanding of whale movement has been derived from whaling records, Discovery marks, photo identification and genetic analyses. However, with advances in satellite tagging technology and concurrent development of analytical methodologies we can now detail finer scale humpback whale movement, infer behavioural context and examine how these animals interact with their physical environment. Here we describe the temporal and spatial characteristics of migration along the east Australian seaboard and into the Southern Ocean by 30 humpback whales satellite tagged over three consecutive austral summers. We characterise the putative Antarctic feeding grounds and identify supplemental foraging within temperate, migratory corridors. We demonstrate that Antarctic foraging habitat is associated with the marginal ice zone, with key predictors of inferred foraging behaviour including distance from the ice edge, ice melt rate and variability in ice concentration two months prior to arrival. We discuss the highly variable ice season within the putative foraging habitat and the implications that this and other environmental factors may have on the continued strong recovery of this humpback whale population.

International audience; Open ocean predator-prey interactions are often difficult to interpret because of a lack of information on prey fields at scales relevant to predator behaviour. Hence, there is strong interest in identifying the biological and physical factors influencing the distribution and abundance of prey species, which may be of broad predictive use for conservation planning and evaluating effects of environmental change. This study focuses on a key Southern Ocean prey species, Antarctic krill Euphausia superba, using acoustic observations of individual swarms (aggregations) from a large-scale survey off East Antarctica. We developed two sets of statistical models describing swarm characteristics, one set using underway survey data for the explanatory variables, and the other using their satellite remotely sensed analogues. While survey data are in situ and contemporaneous with the swarm data, remotely sensed data are all that is available for prediction and inference about prey distribution in other areas or at other times. The fitted models showed that the primary biophysical influences on krill swarm characteristics included daylight (solar elevation/radiation) and proximity to the Antarctic continental slope, but there were also complex relationships with current velocities and gradients. Overall model performance was similar regardless of whether underway or remotely sensed predictors were used. We applied the latter models to generate regional-scale spatial predictions using a 10-yr remotely-sensed time series. This retrospective modelling identified areas off east Antarctica where relatively dense krill swarms were consistently predicted during austral mid-summers, which may underpin key foraging areas for marine predators. Spatiotemporal predictions along Antarctic predator satellite tracks, from independent studies, illustrate the potential for uptake into further quantitative modelling of predator movements and foraging. The approach is widely applicable to other krill-dependent ecosystems, and our findings are relevant to similar efforts examining biophysical linkages elsewhere in the Southern Ocean and beyond.

Body size and feeding mode are two fundamental characteristics that determine foraging performance and ecological niche. As the smallest obligate lunge filter feeders, minke whales represent an ideal system for studying the physical and energetic limits of filter feeding in endotherms. We used multi-sensor suction cup tags to quantify the feeding performance of Antarctic minke whales. Foraging dives around and beneath sea ice contained up to 24 lunges per dive, the highest feeding rates for any lunge-feeding whale. Their small size allows minke whales access to krill in sea-ice environments not easily accessible to larger baleen whales. Furthermore, their ability to filter feed provides an advantage over other smaller sympatric krill predators such as penguins and seals that feed on individual prey. The unique combination of body size, feeding mechanism and sea-ice habitat of Antarctic minke whales defines a previously undocumented energetic niche that is unique among aquatic vertebrates.

Humpback whales undertake long-distance seasonal migrations between low latitude winter breeding grounds and high latitude summer feeding grounds. We report the first in-depth population genetic study of the humpback whales that migrate to separate winter breeding grounds along the northwestern and northeastern coasts of Australia, but overlap on summer feeding grounds around Antarctica. Weak but significant differentiation between eastern and western Australia was detected across ten microsatellite loci (FST = 0.005, P = 0.001; DEST = 0.031, P = 0.001, n = 364) and mitochondrial control region sequences (FST = 0.017 and ΦST = 0.069, P = 0.001, n = 364). Bayesian clustering analyses using microsatellite data could not resolve any population structure unless sampling location was provided as a prior. This study supports the emerging evidence that weak genetic differentiation is characteristic among neighboring Southern Hemisphere humpback whale breeding populations. This may be a consequence of relatively high gene flow facilitated by overlapping summer feeding areas in Antarctic waters.

The overlap and conflict between marine mammals and fisheries is widespread and increasing globally. This study used satellite telemetry to examine the foraging patterns of lactating female New Zealand (NZ) sea lions Phocarctos hookeri from the Auckland Islands (50°50′S, 166°28′E), over two consecutive austral winters, 1996 and 1997. Overall, all individuals tagged were characterised as benthic foragers and were found to be diving beyond their calculated aerobic dive limits on 65% of all dives. They exhibited preferential use of the continental shelf and its edge, as they do in summer (59% overlap between winter and summer foraging areas). This study confirms that NZ sea lion foraging behaviours and areas are similar between seasons, and within and between years. Female winter foraging areas spatially overlapped by 38% with the operation areas of sub-Antarctic trawl fisheries. These findings are important because: female NZ sea lions show extreme foraging behaviour year round; and overlap with tr...

In this study 308 milk samples were collected and analyzed from 181 individual female New Zealand sea lions (NZ sea lions; Phocarctos hookeri) breeding on Enderby Island (Auckland Islands). Samples were collected from the 1st part of early lactation (January and February) over a period of 7 years (1997, 1999–2003, and 2005). The effect of year, month, and maternal characteristics (body mass, body condition index [BCI], and age class) on the composition of milk was evaluated using a mixed model for repeated measures. The gross composition (± SD) of the milk was lipid (21.3% ± 8.1%), protein (9.4% ± 2.4%), water (67.9% ± 8.8%), ash (0.48% ± 0.06%), and energy content (10.3 ± 3.2 kJ/g). Overall, the quality of milk of the NZ sea lions in this study was relatively lower in solids and fats than that of other pinnipeds and, in particular, other sea lion species. There were significant effects of year and month on the concentration of lipids in milk, and of year and maternal age class on maternal body m...

Operational interactions between odontocetes (i.e., toothed whales) and longline gear are a global phenomenon that may threaten the conservation of odontocete populations and the economic viability of longline fisheries. This review attempts to define the issue, summarize the trends and geographical extent of its occurrence over the last half century, explore the potential impact on odontocetes and on fisheries, and describe potential acoustic and physical mitigation solutions. Reports of odontocete bycatch rates are highly variable (between 0.002 and 0.231 individuals killed per set) and at least 20 species may be involved. Information about marine mammal population size, migration patterns and life history characteristics are scarce, although at least one population may be in decline due to losses attributable to longline bycatch. Information about the financial impact of depredation on pelagic longline fisheries is also scarce, although estimates of daily fleet-wide losses range between US$1,034 and US$8,495 (overall fleet income was not reported). Such biological and financial losses may be unsustainable. Recent developments in acoustic and physical mitigation technologies have yielded mixed results. Acoustic mitigation technologies have no moving parts, although require complex electronics. To date, they are insufficiently developed and their efficacy has been difficult to assess. Physical mitigation technologies generally require complex moving parts, although they are relatively simple to develop and assess. Further development and testing remains necessary before widespread implementation would be possible. Development of these approaches should be prioritized and a “toolbox” of various strategies and solutions should be compiled, because a single panacea to the problem is unlikely to emerge.

During the winter months, from June to September, humpback whales Megaptera novaeangliae breed and calve in the waters of the Great Barrier Reef (GBR) after migrating north from Antarctic waters. Clearly defined wintering areas for breeding and calving comparable to those identified in other parts of the world have not yet been identified for humpback whales in the GBR Marine Park (GBRMP), mainly because of its large size, which prohibits broad-scale surveys. To identify important wintering areas in the GBRMP, we developed a predictive spatial habitat model using the Maxent modelling method and presence-only sighting data from nondedicated aerial surveys. The model was further validated using a small independent satellite tag data set of 12 whales migrating north into the GBR. The model identified restricted ranges in water depth (30 to 58 m, highest probability 49 m) and sea surface temperature (21 to 23°C, highest probability 21.8°C) and identified 2 core areas of higher probability of whale occurrence in the GBRMP, which correspond well with the movements of satellite tagged whales. We propose that one of the identified core areas is a potentially important wintering area for humpback whales and the other a migration route. With an estimated increase in port and coastal development and shipping activity in the GBRMP and a rapidly increasing population of whales recovering from whaling off the east Australian coast, the rate of human interactions with whales is likely to increase. Identifying important areas for breeding and calving is essential for the future management of human interactions with breeding humpback whales.

The 2008 IUCN (International Union for Conservation of Nature) review of the status of the world’s mammals identified marine mammals (IUCN 2008) as disproportionally threatened and data poor compared to their terrestrial counterparts, and their status was noted as a particular concern (IUCN 2008, Mace et al. 2008, Schipper et al. 2008). The threats faced by marine mammals were identified as being different, with accidental mortality and pollution being dominant threats that superseded habitat loss, which was identified as the principal concern for land mammals. It was suggested that harvesting remained a major threat for half of the marine mammal species in the world. The distribution of marine mammals was described as being concentrated in tropical and temperate coastal platforms and associated with high levels of primary productivity. Threat levels were highest for marine mammals living in the North Atlantic, North Pacific and Southeast Asia. Range size was described as generally declining toward both poles. Although these generalities are of course correct for all marine mammals, it must be recognized that the larger number of species of cetaceans compared to other marine mammal groups (85 cetaceans, the polar bear Ursus maritimus, 4 extant sirenians and 36 pinniped species) biases the conclusions in the assessment for marine mammals to such a degree that a separate summary for the pinnipeds of the world is warranted. Additionally, a species-level assessment masks some important conservation concerns for pinnipeds with species classified as Least Concern actually containing threatened subspecies ( e.g., ringed seals). Finally, and perhaps most importantly, climate change issues were not specifically addressed in the vast majority of marine mammal assessments done for the 2008 IUCN review (IUCN 2008). We present here a complete assessment of the status and threats facing pinnipeds at the subspecies level, integrating the expected impacts of changing patterns of threat due to global climate change.

The humpback whale, Megaptera novaeangliae, migrates along coastal routes between high latitude summer feeding areas and low latitude winter breeding areas in both hemispheres. The coastal migrations and frequent surfacing behaviour make it easy to take biopsy samples from its skin and blubber and it is consequently the most intensely studied of the great whales. Microsatellite markers have been developed for population genetics and kinship studies of this species, but these show poor genotyping reproducibility among laboratories and scientists. We describe 45 TaqMan® single nucleotide polymorphism markers for a highly reproducable alternative genotyping system for M. novaeangliae.

Chronological age is a fundamental and yet elusive variable in studies of many wild animals. Telomeres are nucleoprotein structures on the ends of chromosomes that change size throughout the life of many animals and because of this property have been advocated as a means to estimate age. In this review, we assess the existing and potential application of using telomeres for age estimation. We argue that there are conceptual and statistical inconsistencies in previous studies and that the basis for telomere change over time is not well understood and affected by several intrinsic and extrinsic process unrelated to chronological time. Furthermore, these processes are likely to vary spatially and temporally for animal populations. We conclude that the current data suggest telomeres should not be used for age estimation. If telomere-based age estimation is to be used, more work in understanding variability in key processes affecting telomere dynamics and rigorous substantiation via blind testing is needed.

Foragers show adaptive responses to changes within their environment, and such behavioural plasticity can be a significant driving force in speciation. We investigated how lactating Antarctic fur seals, Arctocephalus gazella, adapt their foraging within two contrasting ecosystems. Location and diving data were collected concurrently, between December 2003 and February 2004, from 43 seals at Bird Island, where krill, Euphausia superba, are the main prey, and 39 at Heard Island, where mostly fish are consumed. Seals at Heard Island were shorter and lighter than those at Bird Island and they spent longer at sea, dived more frequently and spent more time in the bottom phase of dives. Generalized additive mixed effects models showed that diving behaviours differed between the islands. Both populations exploited diel vertically migrating prey species but, on average, Heard Island seals dived deeper and exceeded their estimated aerobic dive limits. We propose that the recovery of the Heard Island population may be limited by the relative inaccessibility and scarcity of food, whereas at Bird Island, the presence of abundant krill resources helps sustain extremely high numbers of seals, even with increased intra- and inter-specific competition. Both populations of fur seals appear to be constrained by their physiological limits, in terms of their optimal diving behaviour. However, there does appear to be some flexibility in strategy at the level of trip with animals adjusting their time at sea and foraging effort, in order to maximize the rate of delivery of energy to their pups.

Predictable sources of food underpin lifetime reproductive output in long lived animals. The most important foraging areas of top marine predators are therefore likely to be related to environmental features that enhance productivity in predictable spatial and temporal patterns. Even so, although productive areas within the marine environment are distributed patchily in space and time, most studies assess the relationships between feeding activity and proximate, not long term, environmental characteristics. In addition, individuals within a population may exploit different prey types, and these are often associated with different hydrographic features. Until now, models attempting to associate core foraging areas (CFAs) of marine predators with the environmental characteristics of those areas have not considered the diet of individual animals, despite the influence this could have on these relationships. We used bathymetry and multi-year (n=24) mean sea surface temperature and variability as predictors of CFAs of lactating Antarctic fur seals Arctocephalus gazella at Heard Island. The effect of prey types on the predictability of these models was explored by matching diet and foraging trip data of individual seals (n=40 seals, n=1 trip each). Differences in diet between seals were mirrored by their spatial behaviour. Foraging strategies differed both between and within groups of seals consuming different diets. Long-term environmental parameters were useful for predicting the foraging activity of seals that consumed a single prey type with relatively specific habitat preferences, but not for those that consumed single or multiple prey types associated with more varied habitats. Ignoring individual variation in predator diet probably contributes to the poor performance of foraging habitat models. These findings highlight the importance of incorporating individual specialization in foraging behaviour into ecological models and management of predator populations.

Flexibility in foraging behaviour and diet are characteristics of many penguin species. While these 2 aspects of foraging ecology are presumably tightly coupled, the connection between them is rarely examined directly. Using time-depth recorders and satellite telemetry we documented the foraging behaviour of 43 chick-rearing macaroni penguins Eudyptes chrysolophus at Heard Island. We also compared these behavioural data with prey recovered from the stomach contents of each tracked bird. During guard stage, foraging trips were significantly shorter (average duration = 23 ± 18 h, range = 47 ± 40 km) compared to creche stage (average duration = 115 ± 112 h, range = 169 ± 102 km). Creche stage trips also included significantly more shallow dives that appear to be related to travel (39% of dives

Video taken by Greenpeace of whaling by the Japanese whaling fleet in the Southern Ocean provided a unique opportunity to obtain quantitative data relevant to the welfare aspects of the killing of whales. Catches of 16 individual Antarctic minke whales ( Balaenoptera bonaerensis ) were analysed and in two of these asphyxiation appeared the most likely cause of death. Fewer than one in five whales were killed instantaneously and the average time to death for the remaining whales was around 10 min. The presence of Greenpeace did not result in a reduced accuracy of harpoon shots when compared with previous studies.

Mitochondrial ribosomal DNA is commonly used in DNA-based dietary analyses. In such studies, these sequences are generally assumed to be the only version present in DNA of the organism of interest. However, nuclear pseudogenes that display variable similarity to the mitochondrial versions are common in many taxa. The presence of nuclear pseudogenes that co-amplify with their mitochondrial paralogues can lead to several possible confounding interpretations when applied to estimating animal diet. Here, we investigate the occurrence of nuclear pseudogenes in fecal samples taken from bottlenose dolphins (Tursiops truncatus) that were assayed for prey DNA with a universal primer technique. We found pseudogenes in 13 of 15 samples and 1–5 pseudogene haplotypes per sample representing 5–100% of all amplicons produced. The proportion of amplicons that were pseudogenes and the diversity of prey DNA recovered per sample were highly variable and appear to be related to PCR cycling characteristics. This is a well-sampled system where we can reliably identify the putative pseudogenes and separate them from their mitochondrial paralogues using a number of recommended means. In many other cases, it would be virtually impossible to determine whether a putative prey sequence is actually a pseudogene derived from either the predator or prey DNA. The implications of this for DNA-based dietary studies, in general, are discussed.

The Australian sea lion is a listed threatened species under Australian Commonwealth legislation, based on limited population size and the likelihood of further declines in population abundance. Incidental mortality of this species in the west coast rock lobster fishery was identified as a key threat to the distinct sub-population of about 700 ind. on the west coast of Australia. Reports of incidental mortalities in this fishery from a variety of fishery-dependent sources suggested that a minimum of 4 to 5 mortalities occur every fishing season. All reported captures were in shallow waters (

Hays, Graeme C. et al., It is a golden age for animal movement studies and so an opportune time to assess priorities for future work. We assembled 40 experts to identify key questions in this field, focussing on marine megafauna, which include a broad range of birds, mammals, reptiles, and fish. Research on these taxa has both underpinned many of the recent technical developments and led to fundamental discoveries in the field. We show that the questions have broad applicability to other taxa, including terrestrial animals, flying insects, and swimming invertebrates, and, as such, this exercise provides a useful roadmap for targeted deployments and data syntheses that should advance the field of movement ecology., Workshop funding was granted to M.T., A.M.M.S., and C.M.D. by the UWA Oceans Institute, the Australian Institute of Marine Science, and the Office of Sponsored Research at King Abdullah University of Science and Technology (KAUST).

The diet of free-living pinnipeds is most frequently estimated through identification of otoliths, squid mouth-parts and exoskeletons of prey in scats. This is because, although important prey types may not always be detected, sample collection is non-invasive and analysis is easy. Identification of prey DNA in scats is a nascent approach to determining the diet of marine vertebrates that may overcome some of the limitations of hard part analysis. This is the first study to experimentally compare the utility of genetic scatology for identifying consumption of prey types by seals with the occurrence of morphological remains of prey in scats. The occurrences of DNA and hard part remains of one squid and two fish taxa in scats of captive Arctocephalus seals fed mixed prey diets were compared. Both methods detected ingestion of these taxa 7.5–39.5 h prior to defaecation. Although all test prey had robust hard parts, detecting consumption during this period was 1.4 to 5.8 times more likely using genetic analysis than morphological analysis of scats. Based on frequency of occurrence calculations, neither method provided quantitative descriptions of the known diet. Identification of prey using DNA was not compromised by complexity of the diet; each test taxon was unambiguously detected against a background of a multi-species diet. Our results suggest that where diagnostic hard remains of prey are not well represented in scats, or the sample size is small, genetic scatology provides a valuable addition to morphological scat analysis for identifying the recent diet of free-living seals.

A recent review by Ainley et al. has suggested that recent investigations of the ecological structure and processes of the Southern Ocean have “almost exclusively taken a bottom-up, forcing-by-physical-processes approach relating individual species' population trends to climate change”. We examine this suggestion and conclude that, in fact, there has been considerable research effort into ecosystem interactions over the last 25 years, particularly through research associated with management of the living resources of the Southern Ocean. Future Southern Ocean research will make progress only when integrated studies are planned around well structured hypotheses that incorporate both the physical and biological drivers of ecosystem processes.

The diet of pinnipeds is most commonly inferred from morphologically diagnostic remains of prey in their scats. Although this method can generate quantitative estimates of diet simply, important prey types may not always be detected. DNA-based methods improve detection of prey in scats, but they are not quantitative. While some studies have combined morphological and DNA-based methods, these have only assessed prey that are represented by their hard remains in scats. To overcome this bias, we apply molecular and morphological analyses to the soft and hard portions of faecal samples respectively, to estimate the diet of lactating Antarctic fur seals (Arctocephalus gazella) on Heard Island. The diet of this population is of particular interest because it is expanding rapidly and may rely to some extent on mackerel icefish (Champsocephalus gunnari), which are subject to commercial fisheries. Based on results from morphological analysis and likely important prey types, we tested for DNA remains of C. gunnari, myctophids and squid in faecal samples. The proportion of samples (n = 54) yielding no dietary information was reduced from around 25.9% using either method alone, to 9.3% when combined. Detection of all prey types tested for was notably improved by integrating molecular and morphological data. Data from either method alone would have underestimated the number of animals consuming C. gunnari by around 25.7%. Detection of multiple prey types in samples increased from 9.3% when using morphological analysis only, to 33.3% when using DNA only, to 46.3% when using both methods. Taken in isolation, morphological data inferred that individual seals consume either C. gunnari or myctophids, probably foraging in separate locations characteristic of those prey. Including molecular data demonstrated that while this may be true of some individuals, many other seals consume a mixed diet of at least C. gunnari, myctophids and squid. This new approach of combining DNA-based and morphological analyses of diet samples markedly increased the number of samples yielding dietary information, as well as increasing the amount of information attained from those samples. Our findings illustrate the broad potential of this technique to improve insight into trophic interactions in marine ecosystems.

Southern Ocean waters are highly productive and contain important food resources for many indigenous predators, including humans. Management of these resources has fallen under the regulation of the Convention for the Conservation of Antarctic Marine Living Resources (CCAMLR), which has identified a suite of predators as indicator species for monitoring ecosystem fluctuations, including crabeater seals. For crabeater seals to fulfil this role, however, they must respond predictably to fluctuations in krill distribution and abundance. Here, we investigated the validity of using the diving behaviour of this species as an indicator of krill distribution and abun- dance. We used behavioural data collected from 23 crabeater seals fitted with satellite-linked time- depth recorders off eastern Antarctica to quantify habitat use as a function of the amount of time they spent within geographic regions with varying environmental characteristics. This was then linked with diving behaviour in those regions. By integrating geographic location and diving parameters, we demonstrated that habitat use and foraging behaviour within eastern Antarctic waters fluctuated in response to seasonal and spatial environmental variability. Our attempts to use oceanographic variables to develop models of crabeater seal habitat use and behaviour demonstrated real limitations in inferring behaviour from a simple set of environmental factors, but we identified ocean depth and the proximity to the ice edge as factors influencing seasonal habitat use and diving behaviour. Whilst our understanding of the influences driving crabeater seal distribution has improved as a result of telemetry studies, it would appear premature to infer cross-species patterns in distribution and abun- dance with krill given the low predictive power of models derived in the present study. Furthermore, the dynamic and regionally variable use of pelagic habitat by this widely abundant Antarctic preda- tor has important implications for the estimation of crabeater seal biomass.

We developed a computer program (HeardMap) to receive, manage, visualise and analyse Argos location data and synchronised archival dive data. This program was used to acquire a relative intensity scale of areas of foraging and non-foraging activity of marine vertebrate predator populations around Heard Island in near real time during the austral summer of 2003/2004. Foraging and non-foraging areas were determined by using HeardMap, and trawl and acoustic samples were collected using the research ship Aurora Australis to measure the prey characteristics in those areas. A total of 226 meso-pelagic predators were tagged on Heard Island over a 2-month period. Most of these animals foraged in an area to the east of the island. The use of HeardMap demonstrated that large numbers of predators of various species could be tracked simultaneously and near-real-time estimates of foraging intensity could be derived. The inclusion of dive data in the data display provided a measure of behaviour embedded in the spatial distribution of the animals.

In animal ecology, a question of key interest for aquatic species is how changes in movement behavior are related in the horizontal and vertical dimensions when individuals forage. Alternative theoretical models and inconsistent empirical findings mean that this question remains unresolved. Here we tested expectations by incorporating the vertical dimension (dive information) when predicting switching between movement states ("resident" or "directed") within a state-space model. We integrated telemetry-based tracking and diving data available for four seal species (southern elephant, Weddell, antarctic fur, and crabeater) in East Antarctica. Where possible, we included dive variables derived from the relationships between (1) dive duration and depth (as a measure of effort), and (2) dive duration and the postdive surface interval (as a physiological measure of cost). Our results varied within and across species, but there was a general tendency for the probability of switching into "resident" state to be positively associated with shorter dive durations (for a given depth) and longer postdive surface intervals (for a given dive duration). Our results add to a growing body of literature suggesting that simplistic interpretations of optimal foraging theory based only on horizontal movements do not directly translate into the vertical dimension in dynamic marine environments. Analyses that incorporate at least two dimensions can test more sophisticated models of foraging behavior.

Background A population of humpback whales (Megaptera novaeangliae) spends the austral summer feeding on Antarctic krill (Euphausia superba) along the Western Antarctic Peninsula (WAP). These whales acquire their annual energetic needs during an episodic feeding season in high latitude waters that must sustain long-distance migration and fasting on low-latitude breeding grounds. Antarctic krill are broadly distributed along the continental shelf and nearshore waters during the spring and early summer, and move closer to land during late summer and fall, where they overwinter under the protective and nutritional cover of sea ice. We apply a novel space-time utilization distribution method to test the hypothesis that humpback whale distribution reflects that of krill: spread broadly during summer with increasing proximity to shore and associated embayments during fall. Results Humpback whales instrumented with satellite-linked positional telemetry tags (n = 5), show decreased home range size, amount of area used, and increased proximity to shore over the foraging season. Conclusions This study applies a new method to model the movements of humpback whales in the WAP region throughout the feeding season, and presents a baseline for future observations of the seasonal changes in the movement patterns and foraging behavior of humpback whales (one of several krill-predators affected by climate-driven changes) in the WAP marine ecosystem. As the WAP continues to warm, it is prudent to understand the ecological relationships between sea-ice dependent krill and krill predators, as well as the interactions among recovering populations of krill predators that may be forced into competition for a shared food resource.

We documented immediate, behavioural responses of Indo-Pacific bottlenose dolphins (Tursiops sp.) to experimental vessel approaches in regions of high and low vessel traffic in Shark Bay, Western Australia. Experimental vessel approaches elicited significant changes in the behaviour of targeted dolphins when compared with their behaviour before and after approaches. During approaches, focal dolphin groups became more compact, had higher rates of change in membership and had more erratic speeds and directions of travel. Dolphins in the region of low vessel traffic (control site) had stronger and longer-lasting responses than did dolphins in the region of high vessel traffic (impact site). In the absence of additional information, the moderated behavioural responses of impact-site dolphins probably would be interpreted to mean that long-term vessel activity within a region of tourism had no detrimental effect on resident dolphins. However, another study showed that dolphin-watching tourism in Shark Bay has contributed to a long-term decline in dolphin abundance within the impact site (Bejder et al., in press, Conservation Biology). Those findings suggest that we documented moderated responses not because impact-site dolphins had become habituated to vessels but because those individuals that were sensitive to vessel disturbance left the region before our study began. This reinterpretation of our findings led us to question the traditional premise that short-term behavioural responses are sufficient indicators of impacts of anthropogenic disturbance on wildlife.

Summary 1 Foraging behaviours of the Australian sea lion (Neophoca cinerea) reflect an animal working hard to exploit benthic habitats. Lactating females demonstrate almost continuous diving, maximize bottom time, exhibit elevated field metabolism and frequently exceed their calculated aerobic dive limit. Given that larger animals have disproportionately greater diving capabilities, we wanted to examine how pups and juveniles forage successfully. 2 Time/depth recorders were deployed on pups, juveniles and adult females at Seal Bay Conservation Park, Kangaroo Island, South Australia. Ten different mother/pup pairs were equipped at three stages of development (6, 15 and 23 months) to record the diving behaviours of 51 (nine instruments failed) animals. 3 Dive depth and duration increased with age. However, development was slow. At 6 months, pups demonstrated minimal diving activity and the mean depth for 23-month-old juveniles was only 44 ± 4 m, or 62% of adult mean depth. 4 Although pups and juveniles did not reach adult depths or durations, dive records for young sea lions indicate benthic diving with mean bottom times (2·0 ± 0·2 min) similar to those of females (2·1 ± 0·2 min). This was accomplished by spending higher proportions of each dive and total time at sea on or near the bottom than adults. Immature sea lions also spent a higher percentage of time at sea diving. 5 Juveniles may have to work harder because they are weaned before reaching full diving capability. For benthic foragers, reduced diving ability limits available foraging habitat. Furthermore, as juveniles appear to operate close to their physiological maximum, they would have a difficult time increasing foraging effort in response to reductions in prey. Although benthic prey are less influenced by seasonal fluctuations and oceanographic perturbations than epipelagic prey, demersal fishery trawls may impact juvenile survival by disrupting habitat and removing larger size classes of prey. These issues may be an important factor as to why the Australian sea lion population is currently at risk.

We describe seven group-specific primer pairs that amplify small sections of ribosomal RNA genes suitable for identification of animal groups of major importance as prey items in marine ecosystems. These primer sets allow the isolation of DNA from the target animal groups from mixed pools of DNA, where DNA-based identification using universal primers is unlikely to succeed. The primers are designed for identifying prey in animal diets, but could be used in any situation where these animal groups are to be identified by their DNA. © 2006 Blackwell Publishing Ltd.

Eleven lactating female Weddell seals were immobilised using inhaled isoflurane and oxygen, having initially been sedated using an intramuscular injection of midazolam. The seals were selected from colonies in Long Fjord, East Antarctica. Isoflurane was delivered using a precision, out-of-circle vaporiser in a portable, heated, semi-closed circle system anaesthetic machine. Induction time (time from injection of midazolam to detected maximal effect of midazolam) ranged from 12 min to 29 min. The maximal effect of midazolam was assessed as being either moderate sedation (n=9) or heavy sedation (n=2), and the maximal effect of inhaled isoflurane and oxygen was assessed as being light anaesthesia (n=11). The level of chemical restraint achieved using this combination allowed attachment of heart rate monitoring units and collection of biological samples. Recovery time ranged from 1 min to 11 min. The anaesthetic regime proved a practical, safe and reliable method for the immobilisation of lactating Weddell seals under conditions of low environmental temperature.

Giant squids (Architeuthis sp.) remain mysterious; they have evaded observation and are rarely taken from their deep sea habitat. Information on the diet of Architeuthis is scarce due to the limited number of specimens with morphologically recognizable remains in their digestive tracts. We explored the use of polymerase chain reaction (PCR)-based methods for detection of DNA in the prey remains and amorphous slurry from an Architeuthis gut sample. The DNA region amplified varied in size, allowing separation of fish and squid components. Sequence comparisons identified fish prey as Macruronus novaezelandiae. Isolation of Architeuthis DNA from an ingested tentacle and the presence of chitin fragments indicate cannibalism occurs in giant squid. Denaturing gradient gel electrophoresis was used to screen for less common DNA types, revealing a high frequency of PCR-generated false alleles, but no additional prey species.

We quantified the winter and spring movement patterns and foraging behavior of adult crabeater seals (Lobodon carcinophagus), and the influence of sea ice and bathymetry on their foraging behavior. Thirty-four seals (16 M 18 F) were outfitted with Satellite Relay Data Loggers (SRDLs) in the Marguerite Bay Region of the Antarctic Peninsula (� 671S, 671W) during the austral winters of 2001 and 2002. Tags transmitted position and dive information for between 4 and 174 days. Overall, winter activity patterns differed significantly from previously reported data collected during the summer: seals in this study dived deeper (9270.2 m, range 6–713 m) and longer (5.26 min70.6, range 0.2–23.6 min), hauled out during the night rather than the day, and showed seasonal shifts in foraging patterns consistent with foraging on vertically migrating prey. While these patterns were more pronounced in 2001 than in 2002, there were no strong differences in patterns of habitat use between the 2 years. Some animals made long distance movements (furthest movements 664 km to northeast, 1147 km to southwest), but most seals remained within 300 km of their tagging location. Within the Marguerite Bay/Crystal Sound region, seals appeared to favor foraging locations on the continental shelf within the 50 to 450 m depth range, with a tendency to avoid depths of 600 m or greater. In both years, seals remained deep within the pack ice throughout the winter, and did not move into regions with less ice cover. Seals were more likely to be located in shallow water where the bathymetric gradients were greatest, and in areas of higher

Crabeater seals are abundant pack-ice predators that feed almost exclusively on krill. They have a circumpolar distribution and are generally sighted hauled out on ice floes alone or in pairs. Here we report our observations of a sighting of 150–200 crabeater seals, which were synchronised in their diving and surfacing behaviour, along with a summary of similar observations from western Antarctica of large groups of crabeater seals in synchronous dive cycles. We report on the low frequency of sightings of such groups during Antarctic pack-ice seal surveys in eastern (Greater) Antarctica. We examine plausible hypotheses to explain these observations, and suggest this behaviour is likely to represent some form of cooperative foraging behaviour, whereby a net advantage in individual energy intake rates is conferred to each seal. Current research on crabeater seal foraging using satellite-linked dive recorders is unlikely to provide sufficiently fine-scale data to examine this hypothesis. Nor will this approach indicate if a seal is foraging with conspecifics. The use of remote or animal-borne camera systems is more likely to provide an insight into fine-scale foraging tactics, as well as the possible, occasional use of cooperative foraging strategies.

The diving behaviour of 15 dugongs (Dugong dugon) was documented using time–depth recorders (TDRs), which logged a total of 39,507 dives. The TDRs were deployed on dugongs caught at three study sites in northern Australia: Shark Bay, the Gulf of Carpentaria and Shoalwater Bay. The average time for which the dive data were collected per dugong was 10.4±1.1 (S.E.) days. Overall, these dugongs spent 47% of their daily activities within 1.5 m of the sea surface and 72% less than 3 m from the sea surface. Their mean maximum dive depth was 4.8±0.4 m (S.E.), mean dive duration was 2.7±0.17 min and the number of dives per hour averaged 11.8±1.2. The maximum dive depth recorded was 20.5 m; the maximum dive time in water >1.5 m deep was 12.3 min. The effects of dugong sex, location (study site), time of day and tidal cycle on diving rates (dives per hour), mean maximum dive depths, durations of dives, and time spent ≤1.5 m from the surface were investigated using weighted split-plot analysis of variance. The dugongs exhibited substantial interindividual variation in all dive parameters. The interaction between location and time of day was significant for diving rates, mean maximum dive depths and time spent within 1.5 m of the surface. In all these cases, there was substantial variation among individuals within locations among times of day. Thus, it was the variation among individuals that dominated all other effects. Dives were categorised into five types based on the shape of the time–depth profile. Of these, 67% of dives were interpreted as feeding dives (square and U-shaped), 8% as exploratory dives (V-shaped), 22% as travelling dives (shallow-erratic) and 3% as shallow resting dives. There was systematic variation in the distribution of dive types among the factors examined. Most of this variation was among individuals, but this differed across both time of day and tidal state. Not surprisingly, there was a positive relationship between dive duration and depth and a negative relationship between the number of dives per hour and the time spent within 1.5 m of the surface after a dive.

Unique DNA sequences are present in all species and can be used as biomarkers for the detection of cells from that species. These DNA sequences can most easily be detected using the polymerase chain reaction (PCR), which allows very small quantities of target DNA sequence to be amplified even when the target is mixed with large amounts of nontarget DNA. PCR amplification of DNA markers that are present in a wide range of species has proven very useful for studies of species diversity in environmental samples. The taxonomic range of species to be identified from environmental samples may often need to be restricted to simplify downstream analyses and to ensure that less abundant sequences are amplified. Group-specific PCR primer sets are one means of specifying the range of taxa that produce an amplicon in a PCR. We have developed a range of group-specific PCR primers for studying the prey diversity found in predator stomach contents and scats. These primers, their design and their application to studying prey diversity and identity in predator diet are described.

We examined population substructure of bottlenose dolphins (Tursiops sp). in Shark Bay, Western Australia, using 10 highly polymorphic microsatellite loci, and mitochondrial DNA (mtDNA). For microsatellite analysis, 302 different animals were sampled from seven localities throughout the bay. Analysis of genetic differentiation between sampling localities showed a significant correlation between the number of migrants (Nm) calculated from FST, RST and private alleles, and distance between localities–a pattern of isolation-by-distance. For mtDNA, 220 individuals from all seven localities were sequenced for a 351 base pair fragment of the control region, resulting in eight haplotypes, with two distinct clusters of haplotypes. Values of FST and (φ)ST for mtDNA yielded statistically significant differences, mostly between localities that were not adjacent to each other, suggesting female gene flow over a scale larger than the sampled localities. We also observed a significant correlation between the number of female migrants calculated from FST and φST and the distance of sampling localities. Our results indicate that dispersal in female dolphins in Shark Bay is more restricted than that of males.

Single-strand conformation polymorphism (SSCP) analysis of the second exon of the DQB gene in the endemic New Zealand sea lion (Phocarctos hookeri) was automated using fluorescently-labelled PCR primers and a capillary sequencer. SSCP mobility profiles (n = 61 individuals) were confirmed by direct sequencing of genomic polymerase chain reaction (PCR) products (n = 39). We found only two alleles defined by changes at a single codon of this apparently functional locus. SSCP at a constant temperature above ambient allowed consistent and rapid genotyping. By comparison, sequence-based genotyping was highly dependent on the threshold used to identify secondary peaks indicative of a heterozygote.