Southern Elephant Seals Mirounga Leonina Research Articles

Natal site fidelity by breeding female southern elephant seals in relation to their history of participation in the winter haulout

Of the four types of terrestrial haulout periods undertaken by southern elephant seals Mirounga leonina, only the purpose of the winter haulout is unknown. Returning to a haulout site from distant pelagic foraging grounds bears significant costs in terms of increased energy expenditure, reduced foraging time and increased exposure to predation; therefore, each haulout must serve a purpose. We examined the hypothesis that the winter haulout serves to maintain familiarity with the natal site, thereby increasing site fidelity. To this end, we analysed a long-term mark-recapture dataset for female southern elephant seals at Marion Island, Southern Ocean. Results indicate that, whereas greater natal site fidelity as primiparous females was associated with recorded presence ashore at the study site during the winter haulout as immatures, this was not the case for multiparous females. Furthermore, recorded presence ashore during both the moult haulouts as immatures, and all haulouts as immatures, irrespective of haulout type, was also associated with increased site fidelity. This suggests that any haulout at the natal island as an immature seal, whether for the moult or winter haulout, assists in maintaining site fidelity. Therefore, while the winter haulout facilitates greater natal site fidelity, whether this is the sole reason for this terrestrial period remains uncertain.

Elephant seal foraging dives do indeed track prey distribution, but temperature influences the distribution of prey: Reply to Boersch-Supan et al. (2012)

ABSTRACT: McIntyre et al. (2011a; Mar Ecol Prog Ser 441:257−272) illustrated a number of rela-tionships between environmental variables and the dive behaviour of satellite-tracked southernelephant seals Mirounga leonina . One of these associations was that seals tended to increase theirdive depths and spend less time at targeted dive depths when swimming in warmer waters.Boersch-Supan et al.’s (2012; Mar Ecol Prog Ser 461:293−298) comment on this study suggests thatthe link described between dive depths and in situ temperature is actually a link between divedepths and prey distribution. We do not dispute this assertion, having discussed this likelihood inMcIntyre et al. (2011a). Boersch-Supan et al. (2012) further provide a number of criticisms, basedpartly on their observations of potential prey distributions within a comparatively small geo-graphic area. We argue that their results are not directly comparable to those presented inMcIntyre et al. (2011a) given the limited spatial overlap of the study areas and sparse, small-scaledataset presented. We further provide replies to technical comments by Boersch-Supan et al.(2012) pertaining to our data analyses.KEY WORDS: Southern elephant seals · Foraging ecology · Climate change · Marion Island · Bio-logging

Depletion of deep marine food patches forces divers to give up early

Many optimal foraging models for diving animals examine strategies that maximize time spent in the foraging zone, assuming that prey acquisition increases linearly with search time. Other models have considered the effect of patch quality and predict a net energetic benefit if dives where no prey is encountered early in the dive are abandoned. For deep divers, however, the energetic benefit of giving up is reduced owing to the elevated energy costs associated with descending to physiologically hostile depths, so patch residence time should be invariant. Others consider an asymptotic gain function where the decision to leave a patch is driven by patch-depletion effects - the marginal value theorem. As predator behaviour is increasingly being used as an index of marine resource density and distribution, it is important to understand the nature of this gain function. We investigated the dive behaviour of the world's deepest-diving seal, the southern elephant seal Mirounga leonina, in response to patch quality. Testing these models has largely been limited to controlled experiments on captive animals. By integrating in situ measurements of the seal's relative lipid content obtained from drift rate data (a measure of foraging success) with area-restricted search behaviour identified from first-passage time analysis, we identified regions of high- and low-quality patches. Dive durations and bottom times were not invariant and did not increase in regions of high quality; rather, both were longer when patches were of relatively low quality. This is consistent with the predictions of the marginal value theorem and provides support for a nonlinear relationship between search time and prey acquisition. We also found higher descent and ascent rates in high-quality patches suggesting that seals minimized travel time to the foraging patch when quality was high; however, this was not achieved by increasing speed or dive angle. Relative body lipid content was an important predictor of dive behaviour. Seals did not schedule their diving to maximize time spent in the foraging zone in higher-quality patches, challenging the widely held view that maximizing time in the foraging zone translates to greater foraging success.

Unmarked individuals in mark-recapture studies: Comparisons of marked and unmarked southern elephant seals at Marion Island

The presence of unmarked individuals is common in mark–recapture study populations; however, their origin and significance in terms of population dynamics remain poorly understood. At Marion Island, southern Indian Ocean, where virtually all southern elephant seal Mirounga leonina pups born annually (1983–2008) were marked in a long-term mark–resight study, large numbers of unmarked seals occur. Unmarked seals originate either from marker (tag) loss or from immigration. We aimed to identify patterns in the occurrence of marked and unmarked individuals that will allude to the possible origin and significance of the untagged component of the population, predicting that tag loss will add untagged seals to mainly adult age categories whereas migrating untagged individuals will be mostly juveniles. We fitted a generalized linear model using the factors month, year and age-class to explain the relative abundance of untagged seals (tag ratio) from 1997 to 2009. Site usage of untagged seals relative to tagged seals was assessed using a binomial test. Untagged seals, predominantly juveniles, were present in the highest proportions relative to tagged seals during the winter haulout (tagged seals/total seals less than 0.3) and the lowest proportion (approximately 0.5) during the female breeding haulout, increasing in relative abundance from 1997 to 2009. Untagged seals were distributed evenly across suitable haulout sites while tagged seals displayed high local site fidelity and occurred in greater numbers at or near large breeding beaches. Untagged seals are considered to be mostly migrant seals that disperse from other islands within the southern Indian Ocean and haul out at Marion Island during non-breeding haulouts in particular. Some of these seals immigrate to the breeding population, which can be a key component of the local population dynamics. We emphasize the need for mark–recapture studies to evaluate the role of the unmarked component of a population, thereby inducing a more confident estimation of demographic parameters from the marked sample.

Stable carbon and nitrogen isotope variations in canine dentine growth layers of Kerguelen southern elephant seals

Foraging behaviour of mammals, namely the change in distribution and trophic lev- els from juvenile stage to adulthood, can be investigated by measuring δ 13 C and δ 15 N stable iso- topes for layers deposited in a growing tooth. For the first time, we describe geographic differ- ences in the ontogeny of foraging strategies and in the niche partitioning process according to sex and age of a highly sexually dimorphic species: the southern elephant seal Mirounga leonina. Canines from 8 males and 6 females were analysed for δ 13 C and δ 15 N stable isotope signatures. To assess intra-individual variability, instead of analysing collagen we analysed the bulk dentine within each of the 4 growth layers deposited annually. The δ 13 C signature revealed that, in indi- viduals of 1 to 4 yr of age, teeth of both males and females exhibited large intra-individual varia- tion in δ 13 C, suggesting that juveniles were foraging over a broad range of marine habitats encom- passing both sub-Antarctic and Antarctic waters. Four out of the 6 teeth taken from females were collected on individuals younger than 4 yr, preventing investigation of longer-term changes. A δ 13 C pattern emerged for males older than 4 yr: individuals became resident to either a sub- Antarctic (-17‰) or an Antarctic (-20‰, both values reported as deviations from the Vienna PeeDee Belemnite standard) foraging habitat, with a decrease in intra-individual variability. Up to the age of 4 yr, juvenile males were at a slightly higher trophic level than juvenile females, but by the age of 4 yr, while their δ 13 C signature revealed that they were faithful to their foraging habitat, males exhibited a significant increase in their trophic levels, as shown by their δ 15 N signature.

Mesoscale eddies as foraging area of a deep-diving predator, the southern elephant seal

Physical processes, such as mesoscale eddies, play an important role in ocean mixing and concentrating biological productivity, which provides great feeding opportunities for many marine top predators. This study describes the presumed foraging behaviour of southern elephant seals Mirounga leonina within an intense eddy field along the Polar Front in relation to the physical features of the eddies targeted by foraging seals, especially in the vertical dimension. Combining satellite measurements and data from tags deployed on elephant seals, we showed that some seals (10 of 38) preferentially selected the edge of cyclonic structures, generally associated with a high eddy kinetic energy, during intensive foraging phases. Moreover, we observed that elephant seals dived deeper within cyclonic eddies compared with anti-cyclonic structures. The behaviour of seals well illustrates the function of the spin-up of cyclonic eddies, where the upwelling of nutrients occurs at the centre, and high productivity and high concentration of prey occur at the edge. Productivity, aggregation of resources and accessibility of prey associated with cyclonic eddies probably act together to determine the selection of these structures by foraging elephant seals. However, eddies are not the only favourable foraging habitat within the Polar Frontal Zone, and the identification of oceanographic conditions associated with these other foraging zones requires further work.

Looking at the unseen: combining animal bio‐logging and stable isotopes to reveal a shift in the ecological niche of a deep diving predator

Understanding how marine top predators exploit their environment is a central topic in marine ecology. Among all methodologies used to investigate this part of ecology, electronic devices are very useful to track animals' movements and foraging habitats, but they do not provide any dietary information. Stable isotopes provide information on trophic levels but remain imprecise to identify small spatial‐scale habitats. In this study, we combined the two approaches to obtain a synoptic view of the foraging behaviour variability of southern elephant seals Mirounga leonina. Our results suggested marked differences in distribution, diving behaviour, foraging habitats, trophic levels, and dietary habits of elephant seals according to their sex and age. Thus, we characterized main foraging habitats over the Kerguelen‐Heard Plateau and the Antarctic shelf for juvenile males, while females foraged mainly in oceanic waters of the Polar Frontal Zone and the Antarctic Zone. In addition, we highlighted the ontogeny of niche partitioning in this sexually dimorphic species. While females did not exhibit a major dietary shift in relation to their age and their breeding status, a different picture emerged for males. Young males had a trophic level identical to that of all females. However, at 3–4 yr of age, males showed a progressive increase in trophic level. The inter‐annual combination of bio‐logging and stable isotopes could provide a powerful tool to investigate possible shifts in ecological niche between years according to environmental changes.

Southern elephant seals ( Mirounga leonina) along the Brazilian coast: review and additional records

Southern elephant seal (Mirounga leonina) is a huge marine predator that has a widely circumpolar distribution and breeding sites near the subantarctic convergence. Nevertheless, this seal has been found in several locations far from their breeding grounds. In Brazilian waters this species has been occasionally recorded. The aim of this paper is to review the occurrence of this species along the Brazilian coast, adding new records in the territory. From 1954 to 2008 we compiled 46 records of M. leonina distributed in eight Brazilian coastal states. We present eight non-published records which represent 17.4% of all the records for the Brazilian coast. From 32 sexed seals, 81.2% were males and 61.3% of the specimens were immature. Most records occurred during autumn (N = 15) and winter (N = 18) seasons. Unexpectedly, the records did not show a clear pattern of decrease from high to low latitudinal coastal Brazilian states. The nectonic pattern of M. leonina with the lack of physical barriers and currents moving northwards could be the reason for the dispersion of this seal to different areas along South American coast.

Two records of male southern elephant seals (Mirounga leonina) on the Atlantic coast of Uruguay

Two records of male southern elephant seals (Mirounga leonina) on the Atlantic coast of Uruguay

Blubber fatty acid profiles indicate dietary resource partitioning between adult and juvenile southern elephant seals

When resources are limited or patchy, a species may develop some degree of resource partitioning to reduce intra-specific competition. Development of intra-specific resource partitioning is more pronounced in species with clear phenotypic variation among individuals (e.g. age or sex). Southern elephant seals Mirounga leonina have pronounced sexual dimorphism and range widely in size and foraging range between juvenile and adult stages. However, hypothesized diet-based resource partitioning has been less clear due to difficulties in sampling diet while seals are away from breeding islands. We analysed fatty acids (FAs) from blubber of 122 juvenile seals and compared them to FA profiles from blubber of 52 adult females, and to FA profiles from 51 prey species (grouped as fish and squid) to examine evidence for diet-based resource partitioning in the seals. FA signature analysis revealed physiological and dietary differences between ages. Principle components of the 21 FAs from seal blubber and prey parts distinguished prey from seals, and clearly separated prey species into fish and squid classes. FA profiles from adult females differed to those from juveniles, with the former more 'squid-like' and the latter more 'fish-like'. Variation in FA profiles of seals was also apparent between sexes and during different seasons. Differences in diet between juveniles and adult females suggest resource partitioning occurs in response to large metabolic and physiological differences with age that limit juvenile dispersal and diving abilities. By consuming a different suite of prey species relative to adult females, juvenile southern elephant seals may reduce intra-specific competition.

Immobilization of adult male southern elephant seals (Mirounga leonina) during the breeding and molting periods using a tiletamine/zolazepam mixture and ketamine

Seventy-seven immobilizations were carried out on adult male southern elephant seals at Stranger Point, Isla 25 de Mayo (King George Island) using a combination of Zoletil® (tiletamine and zolazepam) and ketamine in order to obtain biological samples. During 2006/2007, 22 males were immobilized at the beginning of their breeding period (EB), 19 of which were recaptured at the end of breeding (LB). Four were given only once at an unknown stage of breeding (USB) and 18 males were immobilized at the beginning of molting (BM). During 2007/2008, 14 adult males were immobilized at an USB. Zoletil® was administered using an automatic discharge device, whereas ketamine was injected directly with a syringe, and was used only when the initial sedation was not enough to carry out the programmed sampling. The initial mean dose of Zoletil® was 1,387 ± 304 mg, which represented 0.60 ± 0.14 mg/kg, range 0.36–1.05, n = 77. In 47 procedures, an average dose of 1.04 ± 0.66 mg/kg of ketamine was added. Mean immobilization time was 34 ± 14 min. In 25 out of the 77 procedures, males showed apnea, which lasted 8 ± 4 min (range 2–15 min). The necessary doses of Zoletil® and ketamine to attain immobilization differed between stages. For animals taken twice, doses (mg/kg) of Zoletil® and ketamine were significantly higher at the beginning than at the end of breeding. During molting, the doses of Zoletil® given were significantly lower than those used during breeding, although the proportion of animals that required ketamine during molting was significantly higher than during breeding. Zoletil® proved to be a safe immobilizing agent for field work on adult males of this species, given the wide range of doses used without any serious consequences. Furthermore, the addition of ketamine was useful when the initial sedation was not satisfactory or for prolonging the immobilization period in a practical and reliable way.

Ecological repercussions of historical fish extraction from the Southern Ocean

AbstractA major mid‐1980s shift in ecological structure of significant portions of the Southern Ocean was partially due to the serial depletion of fish by intensive industrial fishing, rather than solely to climate factors as previously hypothesized. Over a brief period (1969–1973), several finfish stocks were on average reduced to <50%, and finally (mid‐1980s) to <20%, of original size. Despite management actions, few stocks have recovered and some are still declining. Most affected species exhibit K‐selected life‐history patterns, and before exploitation presumably fluctuated in accordance with infrequent strong year classes, as is true of such fish elsewhere. A climate regime, the Southern Annular Mode, once oscillated between two states, but has remained in its ‘positive mode’ since the time of the fish extraction. This may have increased finfish vulnerability to exploitation. As breeding stocks decreased, we hypothesize that availability of annually produced juvenile fish fed upon by upper‐level predators remained low. Correlations between predator populations and fish biomass in predator foraging areas indicate that southern elephant seal Mirounga leonina, Antarctic fur seal Arctocephalus gazella, gentoo penguin Pygoscelis papua, macaroni penguin Eudyptes chrysolphus and ‘imperial’ shag Phalacrocorax spp. – all feeding extensively on these fish, and monitored at Marion, Crozet, Kerguelen, Heard, South Georgia, South Orkney and South Shetland Islands, where fishing was concentrated – declined simultaneously during the two periods of heavy fishing. These patterns indicate the past importance of demersal fish as prey in Antarctic marine systems, but determining these interactions’ ecological mechanisms may now be impossible.

How to weigh an elephant seal with one finger: a simple three-dimensional photogrammetric application

Several studies have developed photogrammetric techniques for indirect mass estima- tion of seals. Unfortunately, these techniques are often narrowly delineated for specific field scenar- ios or species. Many require sophisticated, custom-designed equipment or analytical tools, limiting their applicability. We aimed to devise a photogrammetric technique for accurate volume/mass esti- mation of seals under a variety of field scenarios without manipulation of the animal and with mini- mal equipment. We used Photomodeler Pro 3-dimensional modelling software to estimate the mass of 53 weighed southern elephant seals Mirounga leonina. The method is centred on animal volume estimation in relation to the 3-dimensional area around it, rather than features of the animal itself, an approach that liberates limitations associated with earlier studies. No morphometric body measures are required for such volume/mass estimation. We offer predictive equations that allow high confi- dence in mass estimates relative to measured mass (95% confidence interval of mean deviation from measured mass is from ±1.34 to ±3.83% depending on the field scenario). A single photographer with a measuring stick and non-customised digital photographic equipment can use this technique to determine the mass of an elephant seal anywhere in the field with the push of a button.

Tracking changes in relative body composition of southern elephant seals using swim speed data

Changes in buoyancy during an animal’s time at sea are a powerful tool for inferring spatial and temporal foraging success. Buoyancy can be difficult to measure, but in some species of seal, drift components of dives can be used. We used swim speed data from adult female southern elephant seals Mirounga leonina using geo-locating velocity-time-depth recorders during 2004 postlactation (PL; n = 7) and 2002, 2004 and 2005 post-moult (PM; n = 18) foraging trips to detect periods of passive drifting during diving. In addition to the characteristic drift dives of elephant seals, drifting also occurred during putative foraging dives. We used generalised linear models (GLMs) to examine the relationship between body lipid content measured on land and several diving variables collected within a week of these measurements being taken. The strongest support (deviance explained = 90%) was for the model including drift rate (77%), seal length (12%) and descent rate (2%). Estimates of body lipid, based on the GLM, were predicted for each day of the foraging trips. Areas where seals increased their relative lipid content from one day to the next corresponded well with areas in which the seals spent the greatest amount of time. Inferring foraging success from positive changes in drift rate has so far been limited to elephant seals which perform characteristic drift dives, but the addition of swim speed data to detect short periods of stationary behaviour allows for this method to be expanded to a greater range of ocean predators.

Dispersion during the moult haulout of southern elephant seals at the Courbet Peninsula, Iles Kerguelen

We studied the dispersion of 4-year-old southern elephant seals (Mirounga leonina) along a 75.5 km coastal area at the Courbet Peninsula, Iles Kerguelen, relative to their birth site when they were ashore to moult in early 1984. The seals were mostly faithful to their natal sites, but availability of suitable moulting habitat (e.g. wallows, vegetated areas) influenced seal dispersion. As moult progressed, the seals moved farther away from their initial moult sites and natal sites, but remained largely on the easterly beaches of the Courbet Peninsula. This behaviour would facilitate mark-recapture estimates of age and sex specific survival.

Australian exploration of Heard Island, 1947–1971

In 1947 knowledge of Heard Island was confined to a rough mapping compiled by nineteenth-century sealers, and the results of four scientific expeditions that had briefly investigated the Atlas Cove area. Exploration continued in two distinct periods between 1947 and 1971. In the first period the Australian National Antarctic Research Expeditions (ANARE) built a scientific station at Atlas Cove in 1947, and occupied it continuously until 1955 as an ‘A Class’ meteorological station, a seismic and magnetic observatory, and a base for other scientific studies and for exploration of the island. In the second period four summer expeditions and one wintering expedition worked on the island between 1963 and 1971. The summer expeditions were an ANARE expedition in 1963, an Australian private expedition (The South Indian Ocean Expedition to Heard Island) in 1965, and ANARE expeditions in 1969 and 1971 associated with United States and French expeditions. A United States expedition wintered in 1969. There were no further expeditions until 1980. The years 1947–1971 saw many achievements. Expedition members recorded seven years of synoptic meteorological observations and four years of seismic and magnetic observations. They developed empirical techniques of work, travel, and survival that shaped the collective character of ANARE and were later applied in Antarctica. Despite difficult terrain and consistently bad weather, and the accidental deaths of two men in 1952, unsupported field parties of two or three men travelling on foot explored and mapped in detail the heavily glaciated island, and documented its topography, geology, glaciology and biology. They made three overland circuits of the island, the first ascent of Big Ben (2745 m), and the first recorded landing on the nearby McDonald Islands. Expedition members bred and trained dog teams for later use in Antarctica. They reported the commencement and subsequent progress of massive glacier retreat caused by regional warming, and of the island's colonisation by king penguins (Aptenodytes patagonicus) and antarctic fur seals (Arctocephalus gazella). They also reported measurements of glacier flow and thickness, the palaeomagnetism of Heard Island rocks, behavioural and population studies of southern giant petrels (Macronectes giganteus) and other birds, studies of southern elephant seals (Mirounga leonina) and leopard seals (Hydrurga leptonyx), and the cold stress and acclimatisation experienced by humans working in the island's wet-cold climate. In addition, Heard Island served as a testing ground for men, equipment, scientific programmes, huskies, general administration, and logistics, without which Mawson station could not have been established as successfully as it was in 1954. The American wintering expedition and the French summer expedition contributed to major international geodetic and geophysical investigations. In sum, the expeditions between 1947 and 1971 added much to our knowledge of Heard Island, and they laid down a solid foundation for the work of later expeditions.

The importance of land-based prey for sympatrically breeding giant petrels at sub-Antarctic Marion Island

Northern (Macronectes halli) and southern (M. giganteus) giant petrels breed at different times at sub-Antarctic Marion Island. Long-term census and breeding success data are used to test for competitive overlap between the two species by correlating population trends with those of land-based prey/carrion species. No parameter was singularly important in population regulation of either giant petrel species and the assumed dependence of breeding northern giant petrels on southern elephant seal Mirounga leonina carrion is not entirely supported.

Heavy metals in molted fur of the southern elephant seal Mirounga leonina

Heavy metals in molted fur of the southern elephant seal Mirounga leonina

The dynamics of male harem dominance in southern elephant seals (Mirounga leonina) at the South Shetland Islands

Breeding chronology, harem structure and changes in male harem dominance were studied at Stranger Point, Isla 25 de Mayo/King George Island, principally by extensive field census work during the 2003 breeding season. Males were individually identified and their size estimated by using a photogrammetric method. Peak female haul out for the population occurred on 31 October, when a total of 276 females were observed along 7 km of coastline, distributed in ten harems with a median size of 16 females. Overall sex ratio and harem sex ratio for the breeding population were 1:6.7 and 1:10.6, respectively. A total of 33 males were identified associated with harems. Male size conferred an advantage in terms of dominance hierarchy, since dominant males (4.91±0.15 m) were significantly longer than subordinate males (4.63±0.19 m). Harems were dominated by an average of 4.5 (range 2–7) different males during the breeding season. Elephant seals at Stranger Point breed in very low density aggregations. The main breeding events in this population occurred later than at other breeding sites, which agrees with previous observations in the area. Male movement among harems suggests that differences in mating success among males could be achieved through their different behaviours.

Behaviour and breeding success of gentoo penguins Pygoscelis papua in areas of low and high human activity

A key factor influencing wildlife responses to human activity is the degree to which animals have been previously exposed to human stimuli. On subantarctic Macquarie Island, gentoo penguins Pygoscelis papua breed in areas of high and low human activity (on and off-station, respectively). We investigated the behaviour and breeding success of gentoo penguins on and off-station, by a) comparing the behavioural responses of guarding gentoos before, during and after exposure to standardised pedestrian approaches, and b) employing an observational study to determine how human activity may have contributed to within-season breeding success in light of other environmental and site variables. Behavioural responses to pedestrian visitation by gentoos off-station were significantly stronger than those of birds breeding on-station. However, no relationship was found between pedestrian activity and breeding success off-station. Breeding success was, however, positively related to colony size, and negatively related to the activity of other penguins, the number of nearby southern elephant seal Mirounga leonina harems and the location of colonies within short grassland. On-station, breeding success was amongst the highest recorded for that season. Habituation, predator exclusion and the relevance of these findings for management are discussed.