Variation in body condition during the post-moult foraging trip of southern elephant seals and its consequences on diving behaviour

Mature female southern elephant seals (Mirounga leonina) come ashore only in October to breed and in January to moult, spending the rest of the year foraging at sea. Mature females may lose as much as 50% of their body mass, mostly in lipid stores, during the breeding season due to fasting and lactation. When departing to sea, post-breeding females are negatively buoyant, and the relative change in body condition (i.e. density) during the foraging trip has previously been assessed by monitoring the descent rate during drift dives. However, relatively few drift dives are performed, resulting in low resolution of the temporal reconstruction of body condition change. In this study, six post-breeding females were equipped with time-depth recorders and accelerometers to investigate whether changes in active swimming effort and speed could be used as an alternative method of monitoring density variations throughout the foraging trip. In addition, we assessed the consequences of density change on the swimming efforts of individuals while diving and investigated the effects on dive duration. Both descent swimming speed and ascent swimming effort were found to be strongly correlated to descent rate during drift dives, enabling the fine-scale monitoring of seal density change over the whole trip. Negatively buoyant seals minimized swimming effort during descents, gliding down at slower speeds, and reduced their ascent swimming effort to maintain a nearly constant swimming speed as their buoyancy increased. One per cent of seal density variation over time was found to induce a 20% variation in swimming effort during dives with direct consequences on dive duration.

Immobilisation dose rates and photogrammetry based mass calculations for southern elephant seals during expedition JUB2010

Immobilization and anaesthesia of adult male southern elephant seals (Mirounga leonina) is potentially risky for animals and scientists. A tiletamine/zolazepam injection is considered the most appropriate drug combination for field application in this species. Since appropriate dosages are difficult to assess due to uncertainties in weight estimation, we used photogrammetry-derived weight estimates to ensure precise post hoc calculations of dosages. We report on 15 intramuscular tiletamine/zolazepam immobilizations of post-moult males of the upper weight class at King George Island/Isla 25 de Mayo, in April 2010. Initial injections were made using blowpipe syringes. Mean tiletamine/zolazepam combined dosages of 0.71 mg/kg (SD ± 0.16) ranged between 0.46 and 1.01 mg/kg. In four cases, ketamine was added in dosages between 0.96 and 2.61 mg/kg. Mean induction period was 23 min (± 15), and the mean duration of the procedures from first injection to release of the animals required 96 min (± 51). Four seals exhibited periods of apnoea, and one case of an extended, repetitive, and potentially critical apnoea (> 25 and 8 min) required intervention in order to successfully re-initiate spontaneous respiration. All procedures resulted in proper immobilizations allowing for the deployment of the satellite tags on the seals' heads. The fact that even substantial deviations between the initial weight estimates and the photogrammetry-derived weight estimates had no apparent effect on the course of the immobilization underlines the drugs' wide safety margin in this species.

At surface behaviour of adult male southern elephant seals from King George Island from expedition JUB2000

Adult male southern elephant seals instrumented in 2000 on King George Island (n = 13), travelled both to the north (doi:10.1594/PANGAEA.231580, doi:10.1594/PANGAEA.231585) and to the east (doi:10.1594/PANGAEA.231571, doi:10.1594/PANGAEA.231579, doi:10.1594/PANGAEA.261708, doi:10.1594/PANGAEA.261709, doi:10.1594/PANGAEA.261710, doi:10.1594/PANGAEA.261711) of the Antarctic Peninsula. Five males (doi:10.1594/PANGAEA.231571, doi:10.1594/PANGAEA.231579, doi:10.1594/PANGAEA.231580, doi:10.1594/PANGAEA.261710, doi:10.1594/PANGAEA.231585) remained within 500 km of the island and focusing movements in the Bransfield Strait and around the Antarctic Peninsula. Sea-surface temperatures encountered by these animals showed little variation and they seemed to move about irrespective of sea ice cover, but frequented areas of shallow bathymetry. Three males (doi:10.1594/PANGAEA.261708, doi:10.1594/PANGAEA.261709, doi:10.1594/PANGAEA.261711) moved as far as 75°S to the east of the peninsula, into the Weddell Sea, with maximum distances of more than 1500 km from King George Island. They travelled into the Weddell Sea along the western continental shelf break until they reached the region of the Filchner Trough outflow. Here the bathymetry consists of canyons and ridges which support the intensive mixing between the warm saline waters of the Weddell Gyre and the very cold outflow waters with Ice Shelf water ingredients at the Antarctic Slope Front. Another five data sets were shorter then 40 days, and excluded from analyses (doi:10.1594/PANGAEA.231568, doi:10.1594/PANGAEA.231576, doi:10.1594/PANGAEA.231572, doi:10.1594/PANGAEA.231577, doi:10.1594/PANGAEA.264710). A computer animation was developed to visualize the animal movements in relation to the extent and concentration of sea ice (doi:10.1594/PANGAEA.509404). The need for re-instrumentation of adult males from King George Island is highlighted to investigate whether males continue to travel to similar areas and to obtain higher resolution data.

Dive depth and dive duration data of subadult and adult, male and female southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

Although numerous studies have addressed the migration and dive behaviour of southern elephant seals (Mirounga leonina), questions remain about their habitat use in the marine environment. We report on the vertical use of the water column in the species and the potential lifetime implications for southern elephant seals from Marion Island. Long-term mark-resight data were used to complement vertical habitat use for 35 known individuals tagged with satellite-relay data loggers, resulting in cumulative depth use extrapolated for each individual over its estimated lifespan. Seals spent on average 77.59% of their lives diving at sea, 7.06% at the sea surface, and 15.35% hauled out on land. Some segregation was observed in maximum dive depths and depth use between male and female animals-males evidently being physiologically more capable of exploiting increased depths. Females and males spent 86.98 and 80.89% of their lives at sea, respectively. While at sea, all animals spent more time between 300 and 400 m depth, than any other depth category. Males and females spent comparable percentages of their lifetimes below 100 m depth (males: 65.54%; females: 68.92%), though males spent 8.98% of their lives at depths in excess of 700 m, compared to females' 1.84% at such depths. Adult males often performed benthic dives in excess of 2,000 m, including the deepest known recorded dive of any air-breathing vertebrate (>2,133 m). Our results provide a close approximation of vertical habitat use by southern elephant seals, extrapolated over their lifespans, and we discuss some physiological and developmental implications of their variable depth use.

Dive depth and dive duration data of subadult and adult, male and female southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

Sexual segregation in habitat use occurs in a number of animal species, including southern elephant seals, where differences in migration localities and dive behaviour between sexes have been recorded. Due to the extreme sexual size dimorphism exhibited by southern elephant seals, it is unclear whether observed differences in dive behaviour are due to increased physiological capacity of males, compared to females, or differences in activity budgets and foraging behaviour. Here we use a mixed-effects modelling approach to investigate the effects of sex, size, age and individual variation on a number of dive parameters measured on southern elephant seals from Marion Island. Although individual variation accounted for substantial portions of total model variance for many response variables, differences in maximum and targeted dive depths were always influenced by sex, and only partly by body length. Conversely, dive durations were always influenced by body length, while sex was not identified as a significant influence. These results support hypotheses that physiological capability associated with body size is a limiting factor on dive durations. However, differences in vertical depth use appear to be the result of differences in forage selection between sexes, rather than a by-product of the size dimorphism displayed by this species. This provides further support for resource partitioning and possible avoidance of inter-sexual competition in southern elephant seals.

At surface behaviour data of juvenile southern elephant seals from Marion Island between 2001 and 2006 with links to datasets

Marine mammals forage in dynamic environments characterized by variables that are continuously changing in relation to large-scale oceanographic processes. In the present study, behavioural states of satellite-tagged juvenile southern elephant seals (n = 16) from Marion Island were assessed for each reliable location, using variation in turning angle and speed in a state-space modelling framework. A mixed modelling approach was used to analyse the behavioural response of juvenile southern elephant seals to sea-surface temperature and proximity to frontal and bathymetric features. The findings emphasised the importance of frontal features as potentially rewarding areas for foraging juvenile southern elephant seals and provided further evidence of the importance of the area west of Marion Island for higher trophic-level predators. The importance of bathymetric features during the transit phase of juvenile southern elephant seal migrations indicates the use of these features as possible navigational cues.

Dive depth frequency of 23 adult male southern elephant seals (Mirounga leonina) from Marion Island and King George Island

Access to different environments may lead to inter-population behavioural changes within a species that allow populations to exploit their immediate environments. Elephant seals from Marion Island (MI) and King George Island (KGI) (Isla 25 de Mayo) forage in different oceanic environments and evidently employ different foraging strategies. This study elucidates some of the factors influencing the diving behaviour of male southern elephant seals from these populations tracked between 1999 and 2002. Mixed-effects models were used to determine the influence of bathymetry, population of origin, body length (as a proxy for size) and individual variation on the diving behaviour of adult male elephant seals from the two populations. Males from KGI and MI showed differences in all dive parameters. MI males dived deeper and longer (median: 652.0 m and 34.00 min) than KGI males (median: 359.1 m and 25.50 min). KGI males appeared to forage both benthically and pelagically while MI males in this study rarely reached depths close to the seafloor and appeared to forage pelagically. Model outputs indicate that males from the two populations showed substantial differences in their dive depths, even when foraging in areas of similar water depth. Whereas dive depths were not significantly influenced by the size of the animals, size played a significant role in dive durations, though this was also influenced by the population that elephant seals originated from. This study provides some support for inter-population differences in dive behaviour of male southern elephant seals.

Dive depth and dive duration data of female southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.

Ocean temperature and salinity measured by CTD SRDLs deployed on southern elephant seals from King George Island with links to datasets

West Antarctic ice shelves have thinned dramatically over recent decades. Oceanographic measurements that explore connections between offshore warming and transport across a continental shelf with variable bathymetry toward ice shelves are needed to constrain future changes in melt rates. Six years of seal-acquired observations provide extensive hydrographic coverage in the Bellingshausen Sea, where ship-based measurements are scarce. Warm but modified Circumpolar Deep Water floods the shelf and establishes a cyclonic circulation within the Belgica Trough with flow extending toward the coast along the eastern boundaries and returning to the shelf break along western boundaries. These boundary currents are the primary water mass pathways that carry heat toward the coast and advect ice shelf meltwater offshore. The modified Circumpolar Deep Water and meltwater mixtures shoal and thin as they approach the continental slope before flowing westward at the shelf break, suggesting the presence of the Antarctic Slope Current. Constraining meltwater pathways is a key step in monitoring the stability of the West Antarctic Ice Sheet.

Dive depth and dive duration data of male southern elephant seals from Marion Island between 2004 and 2008 with links to datasets

We describe the habitat use of 22 male southern elephant seals (Mirounga leonina) satellite tagged at Marion Island between 2004 and 2008. While a few areas of increased utilization appeared to be associated with areas of shallower bathymetry (such as sea-floor ridges and fracture zones), seals in our study did not target other areas of shallow bathymetry within close proximity to Marion Island. Rather, most elephant seals foraged pelagically over very deep water where much variation was evident in diel vertical migration strategies. These strategies resulted in generally deeper and longer dives than what has been reported for male elephant seals from other colonies. No significant differences were recorded for dive durations or dive depths between adults and sub-adults. However, younger animals displayed a positive relationship between dive durations and age, as well as between dive depths and age, while these relationships became negative for older animals. Mixed model outputs suggested that seals increased their aerobic fitness as migrations progressed, enabling them to undertake longer dives. We conclude that Marion Island male elephant seals exhibit much variability in dive strategy and are seemingly capable of exploiting a range of different prey types occurring in various depth layers.

At surface behaviour, dive depth and dive duration and associated hydrographic data of adult male southern elephant seals from King George Island from expedition JUB2010 with links to datasets

Understanding the distribution and foraging ecology of major consumers within pelagic systems, specifically in relation to physical parameters, can be important for the management of bentho-pelagic systems undergoing rapid change associated with global climate change and other anthropogenic disturbances such as fishing (i.e., the Antarctic Peninsula and Scotia Sea). We tracked 11 adult male southern elephant seals (Mirounga leonina), during their five-month post-moult foraging migrations from King George Island (Isla 25 de Mayo), northern Antarctic Peninsula, using tags capable of recording and transmitting behavioural data and in situ temperature and salinity data. Seals foraged mostly within the Weddell-Scotia Confluence, while a few foraged along the western Antarctic Peninsula shelf of the Bellingshausen Sea. Mixed model outputs suggest that the at-sea behaviour of seals was associated with a number of environmental parameters, especially seafloor depth, sea-ice concentrations and the temperature structure of the water column. Seals increased dive bottom times and travelled at slower speeds in shallower areas and areas with increased sea-ice concentrations. Changes in dive depth and durations, as well as relative amount of time spent during the bottom phases of dives, were observed in relation to differences in overall temperature gradient, likely as a response to vertical changes in prey distribution associated with temperature stratification in the water column. Our results illustrate the likely complex influences of bathymetry, hydrography and sea ice on the behaviour of male southern elephant seals in a changing environment and highlight the need for region-specific approaches to studying environmental influences on behaviour.