First-year survival of southern elephant seals,Mirounga leonina,at sub-Antarctic Macquarie Island

Juvenile seals branded on the isthmus of Macquarie Island as pups displayed a high degree of philopatry. They returned more often and in greater densities to the northern third of the island within 10 km of their birth sites. Juvenile seals were observed to haul out more frequently and in greater numbers on the east coast as opposed to the west. Juvenile seals typically hauled out on two occasions, once during the winter, and once to moult. The probability of recapturing (resighting) branded and tagged seals was greater during the mid-year haulout. First-year survival estimates were obtained from searches of all Macquarie Island beaches for marked (branded and tagged) seals. From a branded population of 2000 seals, 897 were known to be alive at age 1 year, and minimum first-year survival was calculated at 44.85%. To this minimum estimate was added the number of seals overlooked during systematic and standardised searches of the island, and a revised estimate of 65.60% was calculated. Survival rates calculated using a custom model and a conventional mark-recapture model (MARK) were compared and no differences detected. Actual survival data and probability of sighting estimates were included in the revised estimate of first-year survival of southern elephant seals at Macquarie Island. There were no differences in the number of surviving males and females. Accepted: 25 October 1998     

Cohort and tag-site-specific tag-loss rates in mark–recapture studies: A southern elephant seal cautionary case

Marker-loss is a common feature of mark–recapture studies and important as it may bias parameter estimation. A slight alteration in tag-site of double tagged southern elephant seals (Mirounga leonina), marked at Marion Island from 1983 to 2005 in an ongoing mark–recapture program, had important consequences for tag-loss. We calculated age-specific tag-retention rates and cumulative tag-retention probabilities using a maximum likelihood model selection approach in the software application TAG_LOSS 3.2.0. Under the tag-loss independence assumption, double tag-loss of inner interdigital webbing tags (IIT; 17 cohorts) remained below 1% in the first 5 yr and increased monotonically as seals aged, with higher tag-loss in males. Lifetime cumulative IIT tag-loss was 11.9% for females and 18.4% for males, and equivalent for all cohorts. Changing the tag-site to the outer interdigital webbing (OIT; 6 cohorts) resulted in increased and cohort-dependent tag-loss, although the variation (mean ± 95% CI) in cumulative tag-loss probabilities never exceeded 5.3% between cohorts at similar age. Although different studies may homogenize techniques, we advocate the importance of data set-specific assessment of tag-loss rates to ensure greatest confidence in population parameters obtained from mark–recapture experiments. Permanent marking should be implemented where feasible.     

Shifting trends: detecting environmentally mediated regulation in long-lived marine vertebrates using time-series data

Assessing the status and trends in animal populations is essential for effective species conservation and management practices. However, unless time-series abundance data demonstrate rapid and reliable fluctuations, objective appraisal of directionality of trends is problematic. We adopted a multiple-working hypotheses approach based on information-theoretic and Bayesian multi-model inference to examine the population trends and form of intrinsic regulation demonstrated by a long-lived species, the southern elephant seal. We also determined the evidence for density dependence in 11 other well-studied marine mammal species. (1) We tested the type of population regulation for elephant seals from Marion Island (1986–2004) and from 11 other marine mammal species, and (2) we described the trends and behavior of the 19-year population time series at Marion Island to identify changes in population trends. We contrasted five plausible trend models using information-theoretic and Bayesian-inference estimates of model parsimony. Our analyses identified two distinct phases of population growth for this population with the inflexion occurring in 1998. Thus, the population decreased between 1986 and 1997 (−3.7% per annum) and increased between 1997 and 2004 (1.9% per annum). An index of environmental stochasticity, the Southern Oscillation Index, explained some of the variance in r and N. We determined analytically that there was good evidence for density dependence in the Marion Island population and that density dependence was widespread among marine mammal species (67% of species showed evidence for population regulation). This approach demonstrates the potential functionality of a relatively simple technique that can be applied to short time series to identify the type of regulation, and the uncertainty associated with the phenomenon, operating in populations of large mammals.     

Do southern elephant seals show density dependence in fecundity?

Here we provide an alternative interpretation to that of Pistorius et al. (2001), concerning density-dependent increases in fecundity resulting in population regulation of the southern elephant seal population at Marion Island. We do not contradict the findings of Pistorius et al. (2001), because it does appear: (1) that a change in fecundity has been observed, and (2) that some factor related to food supply is the most likely cause for an observed population decline and increase in reproductive performance. The main observation leading to the interpretation of density-dependent feedback in the population of southern elephant seals at Marion Island (one of the Prince Edward Islands) is that there has been a reduction in the population's rate of decline in recent years (reported by Pistorius et al. (1999b)), and that this could have resulted from a per capita increase in food availability. However, because rates of population change are rarely linearly constant, changes in population size should be expressed on a logarithmic, rather than a linear scale, as used by Pistorius et al. (1999b). Re-plotting the linear values of Pistorius et al. (1999b) on the natural logarithmic scale gave no clear change in the rate of population decline; therefore, we conclude that the rate of population change (decline) has remained constant from 1986 to 1997 (r=-0.0439). The Marion Island population is part of the larger Kerguelen population, and there might be considerable overlap in the foraging areas, and possibly prey, exploited by elephant seals from all sub-populations within this larger population. Changes in the number of intra-specific resource competitors at Marion Island are therefore unlikely to alter per capita food availability since the Marion population constitutes approximately 1% of the total Kerguelen population. We propose an alternative hypothesis that the present data support a mechanism driving the proposed increase in per capita food supply through changes in either: (1) inter-specific food competition, (2) rates of predation, (3) changes in weather pattern or (4) disease.     

Temporal changes in fecundity and age at sexual maturity of southern elephant seals at Marion Island

Our objective was to examine the effect of variation in reproductive parameters on the demography of southern elephant seals at Marion Island. We used age-specific capture probabilities of breeding females in a Cormack-Jolly-Seber context to derive reproductive rates. We found that age at maturity declined and fecundity rates increased as the population declined, indicating a compensatory response. Fecundity rates ranged from 0.03 to 0.29 among 3-year-olds (mean=0.16), 0.18 to 0.50 in 4-year-olds (mean=0.40), and 0.28 to 0.50 in 5-year-olds (mean=0.45). We think that a relative increase in food availability, concomitant with the population decline, promoted earlier sexual maturity correlated with more rapid growth of juveniles when population abundance was lower. It is suggested that the relative importance of fecundity in population regulation in elephant seals has been underestimated. Moreover, it appears that the onset of sexual maturity may be the first demographic variable to change in response to a change in population density.     

Long distance breeding dispersal of a southern elephant seal

Southern elephant seals range extensively during regular foraging excursions. Despite this they are highly philopatric and long range dispersal is rare. At Gough Island, southern Atlantic Ocean, we observed a breeding adult male elephant seal during September 2009, which had been tagged on its natal beach at Marion Island, southern Indian Ocean, in November 1998. The individual was resighted only once on Marion Island, 6 months after tagging. This 3,860 km movement represents dispersal (and likely gene flow) between distinct populations from different elephant seal geographical provinces. Given the polygynous breeding system of this species, the presence of this single male may have a disproportionate genetic effect on the small number of southern elephant seals breeding at Gough Island.     

Population status, trends and a re-examination of the hypotheses explaining the recent declines of the southern elephant seal Mirounga leonina

1. Between the 1950s and 1990s the southern elephant seal Mirounga leonina underwent large decreases in population size throughout most of its breeding range in the Southern Ocean. While current population estimates suggest a recent recovery, some breeding populations have continued to decrease in recent years (Macquarie and Marion Islands), others have either remained stable (South Georgia, Kerguelen and Heard Island) or have increased (Peninsula Valdés, Argentina). 2. Intrinsic hypotheses for patterns of regional decline include factors that are affected by density-dependent mechanisms: (i) paucity of males, (ii) population ‘overshoot’ and (iii) pandemic disease. Extrinsic hypotheses include (iv) predation, (v) competition with fisheries concerns, (vi) interspecific competition, (vii) environmental change and (viii) human disturbance. Of the eight hypotheses proposed and examined here, we conclude that three can be discounted (i, v, viii), three are unlikely, but may require more testing (ii, iii, iv) and two are plausible (vi, vii). 3. The interspecific competition hypothesis is difficult to test because it requires the simultaneous monitoring of species that overlap directly with elephant seals, many of which have not been identified or little is known. However, an analysis of the relationship between log variance and log abundance (Taylor's power law) for populations of southern and northern elephant seals suggests that interspecific competition is not a significant factor in the decline of the southern elephant seal. 4. The hypothesis that decreases in southern elephant seal populations between the 1950s and 1990s were caused by the environmental change is the easiest to test and most plausible of the hypotheses. We propose a framework by which to test this hypothesis to determine how food availability affects individual survival.     

Mummified and skeletal southern elephant seals (Mirounga leonina) from the Victoria Land Coast,Ross Sea,Antarctica

We report on an accumulation of mummified southern elephant seals (Mirounga leonina) from Inexpressible Island on the Victoria Land Coast (VLC), western Ross Sea, Antarctica. This accumulation is unusual, as elephant seals typically breed and molt on sub‐Antarctic islands further north and do not currently occupy the VLC. Prior ancient DNA analyses revealed that these seals were part of a large, Antarctic breeding population that crashed ~1,000 yr ago. Radiocarbon dates for Inexpressible Island mummies range from 380 to 3,270 yr before present, too old to have been created by Scott's Northern Party in 1912 and varying too widely in age to represent a catastrophic death assemblage. Skeletal measurements reveal that most Inexpressible Island mummies are adult or subadult males. The presence of male elephant seals on Inexpressible Island until several hundred years ago suggests that, at a minimum, it served as a haul‐out site for the large Antarctic population and may have hosted a breeding colony. The conditions that allowed this Antarctic population to use the Ross Sea, the factors spurring its decline, and the implications for the adaptability and sensitivity of the species to environmental change all merit further study.     

Leucistic southern elephant seal at Marion Island

We observed a light coloured female southern elephant seal juvenile (Mirounga leonina) twice at Marion Island in August 2008 and confirmed that it was leucistic rather than albinistic. Though there have been a few previous reports of light-coloured southern elephant seals, this is the first confirmed case of leucism in this species. Judged to be 1-year old, perhaps 2-years old at the most, and because we have not observed any leucistic pups at Marion Island during the past 2 years despite an extensive monitoring and tagging program, we think that this animal was born at nearby Prince Edward Island or perhaps further afield at Îles Crozet.     

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.     

The population trend of southern elephant seals (<Emphasis Type="Italic">Mirounga leonina</Emphasis> L.) at Macquarie Island (1952–2004)

Total numbers of adult female southern elephant seals (cows) breeding at Macquarie Island were determined for 19 of the 52 year period between 1952 and 2004. Totals for 1952–1987 (exc. 1959 and 1985) were estimated from the relationship between censuses of the isthmus study area and concurrent censuses for the whole island. Totals for 1987–2004 were obtained by direct census of the entire island in mid-October. Cow numbers decreased from a maximum of about 40,000 in the 1950s to a minimum of 18,300 in 2000, but then increased slightly to 19,200 in 2004. Nonlinear and post-hoc linear analysis of the count data identified 1999 as the year when the exponential rate of change (r) slowed from −1.4% per annum to near zero. The rate of change was not uniform for each census sub-area counted (1987–2004), suggesting that certain terrestrially based density-dependent mechanisms were influencing the annual distribution of cows.     

Pup mortality in southern elephant seals at Marion Island

Pre- and early post-weaning pup mortality of southern elephant seals (Mirounga leonina) at Marion Island from 1990 through 1999 ranged from 1.6% to 7.3% and averaged 3.8%. Mortality of pups after weaning before their first trip to sea accounted for only 12% of the total mortality. We found no relationship between population size and percentage pup mortality, indicating that pup survival is independent of seal density, at least at the densities of breeding seals that prevailed. Indeed, pup mortality was greatest in the smallest harems, apparently owing to a greater number of younger, less experienced mothers. Small harems were generally also found on less suitable beaches than larger harems and this could have contributed to pup injury as a cause of pup mortality on these beaches. Mother-pup separation and injury caused by beachmasters is likely to be responsible for pup mortality in the larger harems. The low rates of pup mortality observed in this study obviate it being a major population regulating agent at Marion Island.     

MOVEMENTS OF SOUTHERN ELEPHANT SEALS AND SUBANTARCTIC FUR SEALS IN RELATION TO MARION ISLAND

Based primarily on an intensive marking/resighting program conducted at Marion Island (46°54'S, 37°45'E) in the Southern Ocean, the inter-island movements of southern elephant seals, Mirounga leonina , and fur seals Arctocephalus spp., were investigated to elucidate their little known pelagic phase. Southern elephant seals, in particular immature animals, readily move between the proximate Marion and Prince Edward islands. Some range as far afield as Iles Crozet, approximately 1,000 km distant where they haul out for the summer molt or during an autumn resting phase. The exchange of individuals between Marion Island and Iles Crozet during the return of immatures for the molt after a winter at sea, suggests overlapping of the foraging ranges of the two populations. Despite their wanderings, the majority of M. leonina from Marion Island probably feed in the proximity of the island, and relocate onto the island for breeding, molting and resting. Of the fur seals, only a few A. tropicalis were seen away from their natal island, in some cases covering distances in excess of 2,000 km. displaying a remarkable dispersal capacity.     

Changes in population sizes and distribution of fur seals at Marion Island

 Population censuses of the Antarctic fur seal (Arctocephalus gazella) and the sub-Antarctic fur seal (A. tropicalis) were conducted during the 1994/1995 breeding season at Marion Island. Pup numbers, determined from direct counts and a mark-recapture experiment, were used to estimate population sizes. Pup numbers of A. tropicalis showed a mean annual change of 2.0% over the previous 6 years, culminating in an estimated total population of 49, 523 for 1994/1995. The population appears to be entering the maturity phase of population growth and may therefore have recovered from the effects of uncontrolled sealing that ended in the early twentieth century. Numbers at the major colonies on Marion Island showed little change since 1989 and these sites may have reached carrying capacity. The extension of breeding to other parts of the island continues. Over the same period, A. gazella pup numbers showed a mean annual change of 17% and the total population numbered 1,205 in 1994/1995. This species has possibly entered the rapid recolonisation phase of population growth. A few hybrid seals were found. Received: 25 October 1995/Accepted: 14 April 1996     

The role of eddies in the diving behaviour of female southern elephant seals

As the Antarctic Circumpolar Current crosses the South-West Indian Ocean Ridge, it creates an extensive eddy field characterised by high sea level anomaly variability. We investigated the diving behaviour of female southern elephant seals from Marion Island during their post-moult migrations in relation to this eddy field in order to determine its role in the animals’ at-sea dispersal. Most seals dived within the region significantly more often than predicted by chance, and these dives were generally shallower and shorter than dives outside the eddy field. Mixed effects models estimated reductions of 44.33 ± 3.00 m (maximum depth) and 6.37 ± 0.10 min (dive duration) as a result of diving within the region, along with low between-seal variability (maximum depth: 5.5 % and dive duration: 8.4 %). U-shaped dives increased in frequency inside the eddy field, whereas W-shaped dives with multiple vertical movements decreased. Results suggest that Marion Island’s adult female elephant seals’ dives are characterised by lowered cost-of-transport when they encounter the eddy field during the start and end of their post-moult migrations. This might result from changes in buoyancy associated with varying body condition upon leaving and returning to the island. Our results do not suggest that the eddy field is a vital foraging ground for Marion Island’s southern elephant seals. However, because seals preferentially travel through this area and likely forage opportunistically while minimising transport costs, we hypothesise that climate-mediated changes in the nature or position of this region may alter the seals’ at-sea dispersal patterns.     

The impact of future summer temperature on public health in Barcelona and Catalonia,Spain

Several epidemiological studies have reported associations between increases in summer temperatures and risks of premature mortality. The quantitative implications of predicted future increases in summer temperature, however, have not been extensively characterized. We have quantified these effects for the four main cities in Catalonia, Spain (Barcelona, Tarragona, Lleida, Girona). We first used case-crossover analysis to estimate the association between temperature and mortality for each of these cities for the period 1983 to 2006. These exposure–response (ER) functions were then combined with local measures of current and projected changes in population, mortality and temperature for the years 2025 and 2050. Predicted daily mean temperatures were based on the A1B greenhouse gas emission, “business-as-usual” scenario simulations derived from the ENSEMBLES project. Several different ER functions were examined and significant associations between temperature and mortality were observed for all four cities. For these four cities, the age-specific piecewise linear model predicts 520 (95%CI  340, 720) additional annual deaths attributable to the change in temperature in 2025 relative to the average from the baseline period of 1960–1990. For 2050, the estimate increases to 1,610 deaths per year during the warm season. For Catalonia as a whole, the point estimates for those two years are 720 and 2,330 deaths per year, respectively, or about 2 and 3% of the warm season. In comparing these predicted impacts with current causes of mortality, they clearly represent significant burdens to public health in Catalonia.     

Body mass loss of moulting female southern elephant seals,Mirounga leonina,at Macquarie Island

Thirteen female southern elephant seals moulting at Macquarie Island lost an average of 4.46±0.80 kg/day (10.01±1.20g/kg/day). There was no significant difference between this rate of body mass loss and that reported for moulting female southern elephant seals from South Georgia. Moulting female southern elephant seals however exhibited larger mass specific mass loss than either female northern elephant seals or male southern elephant seals, indicating a higher metabolic cost of moult in these animals.     

Inter-population differences in diving behaviour of adult male southern elephant seals (Mirounga leonina)

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.     

Penguins, fur seals, and fishing: prey requirements and potential competition in the South Shetland Islands, Antarctica

Antarctic and sub-Antarctic seabirds, marine mammals, and human fisheries concentrate their foraging efforts on a single species, Antarctic krill (Euphausiasuperba). Because these predators may have a significant effect on krill abundance, we estimated the energy and prey requirements of Adelie (Pygoscelisadeliae), chinstrap (Pygoscelisantarctica), and gentoo (Pygoscelispapua) penguins and female Antarctic fur seals (Arctocephalusgazella) breeding on the South Shetland Islands, Antarctica and compared these estimates with catch statistics from the Antarctic krill fishery. Published data on field metabolic rate, population size, diet, prey energy content, and metabolic efficiency were used to estimate prey requirements of these breeding, adult, land-based predators and their dependent offspring. Due to their large population size, chinstrap penguins were the most significant krill predators during the period examined, consuming an estimated 7.8 × 10⁸ kg krill, followed by Adelie penguins (3.1 × 10⁷ kg), gentoo penguins (1.2 × 10⁷ kg), and Antarctic fur seals (3.6 × 10⁶ kg). Total consumption of all land-based predators on the South Shetland Islands was estimated at 8.3 × 10⁸ kg krill. The commercial krill fishery harvest in the South Shetland Island region (1.0 × 10⁸ kg) was approximately 12% of this. Commercial harvest coincides seasonally and spatially with peak penguin and fur seal prey demands, and may affect prey availability to penguins and fur seals. This differs from the conclusions of Ichii et al. who asserted that the potential for competition between South Shetland predators and the commercial krill fishery is low. Received: 26 August 1997 / Accepted: 16 December 1997     

Age-related reproductive variation in a wild marine mammal population

Life history theory predicts a change in reproduction success with age as energy resources are limited and must be allocated effectively to maximize reproduction and survival. In this study, we use three reproductive performance measures, maternal expenditure, offspring weaning mass, and first-year survival, to investigate the role that maternal age plays in successful reproduction. Long-term uninterrupted life history data available for Marion Island’s southern elephant seals and mass change estimates from photogrammetry data allow for assessment of age-related reproduction performance and trade-offs. Known-aged adult females were photographed for photogrammetric mass estimation (n = 29) and their pups weighed at weaning during the 2009 breeding season. Maternal age and proportional mass loss positively influenced pup weaning mass. In turn, first-year pup return rates (as a proxy for survival) were assessed through the intensive mark–recapture program. Pup survival increased with female age and weaning mass. Pups of young females aged 3–6 years have a lower first-year survival probability compared with pups of older and larger females.