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     

COMPARISON OF THE STOMACH CONTENTS OF SOUTHERN ELEPHANT SEALS, MIROUNGA LEONINA, AT MACQUARIE AND HEARD ISLANDS

Abstract: There are three major breeding populations of southern elephant seals centered on Macquarie Island, Kerguelen-Heard Islands and South Georgia-Antarctic Peninsula. The composition of the diet differs between these populations based on published data from Signy Island and data presented here from Macquarie and Heard Islands. These differences in diet appear to be linked to the location at which seals were sampled ranging from the least Antarctic (Macquarie Island) to the most Antarctic (Signy Island). The major food remains consisted of cephalopod beaks and fish eye lenses. More benthic material was found at Heard Island than at Macquarie Island. The diet at Macquarie Island differed between summer and winter and between young animals and adults. The difficulty in collecting dietary samples of southern elephant seals near their main foraging areas makes the study of the feeding ecology of this species extremely difficult in comparison with other Southern Ocean species.     

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.     

Arbovirus of marine mammals: a new alphavirus isolated from the elephant seal louse, Lepidophthirus macrorhini

A novel alphavirus was isolated from the louse Lepidophthirus macrorhini, collected from southern elephant seals, Mirounga leonina, on Macquarie Island, Australia. The virus displayed classic alphavirus ultrastructure and appeared to be serologically different from known Australasian alphaviruses. Nearly all Macquarie Island elephant seals tested had neutralizing antibodies against the virus, but no virus-associated pathology has been identified. Antarctic Division personnel who have worked extensively with elephant seals showed no serological evidence of exposure to the virus. Sequence analysis illustrated that the southern elephant seal (SES) virus segregates with the Semliki Forest group of Australasian alphaviruses. Phylogenetic analysis of known alphaviruses suggests that alphaviruses might be grouped according to their enzootic vertebrate host class. The SES virus represents the first arbovirus of marine mammals and illustrates that alphaviruses can inhabit Antarctica and that alphaviruses can be transmitted by lice.     

Plasma melatonin concentration in neonatal northern elephant seals,Mirounga angustirostris

The development of pineal function in northern elephant seals was examined in an attempt to understand the physiological basis for previously observed high daytime levels of melatonin in neonatal southern elephant seals. Pineal glands from four northern elephant seal pups, estimated age less than 1 week, weighed 3.0 ± 0.80 g, which was significantly less than that previously found in southern elephant seals (4.6 ± 0.35 g). Midday concentrations of plasma melatonin in pups averaged more than 3000 pmol/l in the first 5 days post-partum, but declined rapidly to less than 400pmol/l after day 9. Daytime melatonin levels in northern elephant seals tended to be lower than in southern elephant seals, although they were very high compared with other species. A circadian cycle of plasma melatonin concentration was observed in newborn northern elephant seals, with levels of 3000–5000 pmol/1 during the day, rising to more than 10,000 pmol/1 late in the dark phase. Soon after weaning at 4 weeks of age, daytime and night-time levels were in the range 60–100 pmol/1 and 100–400 pmol/1, respectively. When approximately 10 weeks old, most samples were in the range 100–400 pmol/1 with no discernible difference between day and night levels. The results do not support the hypothesis that the pineal gland is involved in thermogenesis in new-born southern elephant seals. Instead, the very active pineal gland may contribute to energy conservation, by lowering body temperature, particularly at night. As physical insulation is acquired by the deposition of blubber, the mechanism is not required and melatonin falls to adult levels.     

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.     

Mass of weaned elephant seal pups in areas of low and high human presence

On sub-Antarctic Macquarie Island, we examined pup weaning mass of southern elephant seals in relation to human presence. Pup weaning mass was previously found to be positively associated with 1st-year survivorship. Weaned pups were weighed in a remote area, Middle Beach, and in an area of relatively high human presence, Isthmus East. The areas were reasonably similar in beach topography, wind and surf conditions, numbers of seals present per kilometre of coastline, and numbers of males and females present in harems. For a sub-sample of measured pups, data on the respective maternal size were collected using photogrammetry. Both male and female weaned pups on Middle Beach were significantly heavier than those on Isthmus East. Estimated length of mothers was significantly higher on Middle Beach. In proportion to their own size, mothers in both areas produced weaners of similar mass, indicating no direct effect of human disturbance on the efficiency of lactation. It remained unclear whether the area differences in maternal and pup size were due to natural or human-related factors.     

DEMOGRAPHY AND BREEDING BIOLOGY OF A SMALL,LOCALIZED POPULATION OF SOUTHERN ELEPHANT SEALS (MIROUNGA LEONINA)

Southern elephant seals have been studied in depth in most of their breeding range. One notable exception is the Falkland Islands population. We present data on demography and breeding biology of elephant seals of Sea Lion Island, the main breeding site of this species in the Falklands. Sea Lion Island shelters a small, localized population of southern elephant seals (516 breeding females in 1995 and 518 in 1996). Comparison with the few available census data collected prior to our study suggests that the population has been stable in the short term (1989-1996). Females produced pups at maximum rate and pup mortality was low (2.13%). Breeding sex ratio was strongly unbalanced, with about 14 females per breeding male and 47 females per harem-holding male at peak haul-out. Survival rate between breeding seasons was 67.4% for females and 50% for males. Timing of the breeding season was very similar to that recorded in other populations and was in accordance with clinal variation with latitude. Sex ratio at birth was balanced, and no significant weight dimorphism at weaning between sexes was detected (males: 135.4 kg; females: 132.0 kg). Weaning weight was correlated with size class of the mother.     

Bottom-up regulation of a pole-ward migratory predator population

As the effects of regional climate change are most pronounced at polar latitudes, we might expect polar-ward migratory populations to respond as habitat suitability changes. The southern elephant seal (Mirounga leonina L.) is a pole-ward migratory species whose populations have mostly stabilized or increased in the past decade, the one exception being the Macquarie Island population which has decreased continuously over the past 50 years. To explore probable causes of this anomalous trend, we counted breeding female seals annually between 1988 and 2011 in order to relate annual rates of population change (r) to foraging habitat changes that have known connections with atmospheric variability. We found r (i) varied annually from −0.016 to 0.021 over the study period, (ii) was most effected by anomalous atmospheric variability after a 3 year time lag was introduced (R = 0.51) and (iii) was associated with sea-ice duration (SID) within the seals’ foraging range at the same temporal lag. Negative r years may be extrapolated to explain, at least partially, the overall trend in seal abundance at Macquarie Island; specifically, increasing SID within the seals foraging range has a negative influence on their abundance at the island. Evidence is accruing that suggests southern elephant seal populations may respond positively to a reduced sea-ice field.     

Past and present status of the southern elephant seal (Mirounga leonina) at Macquarie Island

The southern elephant seal population at Macquarie Island has undergone a serious decline since regular surveys were commenced in 1949. Approximately 2,900 cows and 250 bulls were counted in the isthmus study area during the 1985 breeding season. Comparisons with 20 other counts made in the same area between 1949 and 1984 showed that, although there was considerable fluctuation between the years, the number of seals ashore has dropped at an average rate of 2.1% per year, resulting in a net decrease of approximately 50% for both males and females. This is similar to the ratio of decline of elephant seal populations in the southern Indian Ocean. The census information from all major elephant seal populations was reviewed and it was concluded that there may be a common factor, or group of factors, acting to reduce the Macquarie Island and Indian Ocean populations, while the populations of the South Atlantic seem to be stable. Several potential explanations for these observations are advanced, but it is concluded that a greater knowledge of the ecology of the seals is needed if the declines are to be understood.     

Fur seals at Macquarie Island: post-sealing colonisation,trends in abundance and hybridisation of three species

Commercial sealers exterminated the original fur seal population at Macquarie Island in the early 1800s. The first breeding record since the sealing era was not reported until March 1955. Three species of fur seal now occur at Macquarie Island, the Antarctic (Arctocephalus gazella), subantarctic (A. tropicalis) and New Zealand (A. forsteri) fur seal. Census data from 54 breeding seasons in the period 1954–2007 were used to estimate population status and growth for each species. Between the 1950s and 1970s, annual increases in pup production for the species aggregate were low. Between 1986 and 2007, pup production of Antarctic fur seals increased by about 8.8% per year and subantarctic fur seals by 6.8% per year. The New Zealand fur seal, although the most numerous fur seal species on Macquarie Island, has yet to establish a breeding population, due to the absence of reproductively mature females. Hybridisation among species is significant, but appears to be declining. The slow establishment and growth of fur seal populations on Macquarie Island appears to have been affected by its distance from major population centres and hence low immigration rates, asynchronous colonisation times of males and females of each species, and extensive hybridisation.     

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.     

The effects of El Niño–La Niña on reproductive parameters of elephant seals feeding in the Bellingshausen Sea

Aim To assess the impacts of El Niño–La Niña events on the pup weaning mass and diet of female southern elephant seals (Mirounga leonina) feeding in the Bellingshausen Sea, Antarctica, and to understand the ecological processes that drive these impacts. Location Atlantic southern elephant seal weaning mass and diet were measured at King George Island (62º14′ S, 58º30′ W). Feeding areas for pregnant female seals from King George Island are located west of Alexander Island in the Bellingshausen Sea. Methods Data on weaning mass were collected between 1985 and 1994 during the breeding season (September–November). Moulting females were anaesthetized and cephalopod beaks were isolated and identified from stomach contents obtained from stomach lavages. Sea‐surface temperature anomaly (SSTA) data for the ‘El Niño 3.4’ geographical region (5º N–5º S, 120º W–170º W) were used to define El Niño–Southern Oscillation (ENSO) event years (grouped as El Niño, La Niña and Neutral) as well as the strength of each ENSO event year. Using data from the US National Center for Environmental Prediction, temperature, sea ice concentration and atmospheric pressure anomalies in the Bellingshausen Sea were calculated from March to August, corresponding to the feeding period of pregnant female seals. Results Positive temperature anomalies and negative pressure anomalies in the Bellingshausen Sea were observed during La Niña years and negative temperature anomalies and positive pressure anomalies during El Niño years. These data correlate with sea ice concentration anomalies, which are highly negative during La Niña years and highly positive during El Niño years. Warm temperature conditions in the Bellingshausen Sea during La Niña years are strongly related to both higher weaning mass in elephant seals and to an increase in squid beaks in the stomach contents of females. Main conclusions It is possible that higher elephant seal weaning masses in La Niña years correlate with warmer waters in the Bellingshausen Sea leading to the rapid growth of squid and their more frequent descents to depths frequented by elephant seals. This results in increased predation by pregnant females, leading to a greater mass among weaned pups. This hypothesis may guide future research about interactions between climate and the marine biosphere.     

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.     

Effects of environmental variables on surface temperature of breeding adult female northern elephant seals,Mirounga angustirostris,and pups

Pinnipeds spend extended periods of time on shore during breeding, and some temperate species retreat to the water if exposed to high ambient temperatures. However, female northern elephant seals (Mirounga angustirostris) with pups generally avoid the water, presumably to minimize risks to pups or male harassment. Little is known about how ambient temperature affects thermoregulation of well insulated females while on shore. We used a thermographic camera to measure surface temperature (T_s) of 100 adult female elephant seals and their pups during the breeding season at Point Reyes National Seashore, yielding 782 thermograms. Environmental variables were measured by an onsite weather station. Environmental variables, especially solar radiation and ambient temperature, were the main determinants of mean and maximum T_s of both females and pups. An average of 16% of the visible surface of both females and pups was used as thermal windows to facilitate heat loss and, for pups, this area increased with solar radiation. Thermal window area of females increased with mean T_s until approximately 26 °C and then declined. The T_s of both age classes were warmer than ambient temperature and had a large thermal gradient with the environment (female mean 11.2±0.2 °C; pup mean 14.2±0.2 °C). This large gradient suggests that circulatory adjustments to bypass blubber layers were sufficient to allow seals to dissipate heat under most environmental conditions. We observed the previously undescribed behavior of females and pups in the water and determined that solar radiation affected this behavior. This may have been possible due to the calm waters at the study site, which reduced the risk of neonates drowning. These results may predict important breeding habitat features for elephant seals as solar radiation and ambient temperatures change in response to changing climate.     

THE ROLE OF PREDATION IN THE ECOLOGY OF THE RINGED SEAL IN BARROW STRAIT, NORTHWEST TERRITORIES, CANADA

Predation on ringed seals ( Phoca hispida ) was examined in Barrow Strait between March and May 1984 to 1986. Polar bears were the most important predator. Evidence of bear predation was observed at 18–30% of the ringed seal subnivean structures we located. Ten to 24% of predation attempts were successful, with pups making up 75% to 100% of the seals killed. Bears killed an average of 0.08 to 0.51 seals/km², which comprised 8 to 44% of the estimated annual pup production. Bears were successful on average in 11.3% of their attempts to kill pups hidden inside birth lairs. On southeast Baffin Island where snow was soft and pups were exposed, bears were successful in 33.5% of their attempts to kill a seal. Negative correlations were found between mean snow depth and predation by polar bears ( r = -0.896, P = 0.04, n = 5) in 1985, and between snow depth and the number of predation attempts ( r = -0.613, P = 0.02, n = 14) in 1986.     

Counts of southern elephant seals, Mirounga leonina, at Bouvet Island

Southern elephant seals were counted and classified into subjective sex-age classes on a weekly basis during expeditions to Bouvet Island in the austral summers of 1996/1997 and 1998/1999. The expeditions coincided with the moulting period of elephant seals aged one?year and older. The presence of weaned pups at the principal haulout site, Nyrøysa/Westwindstranda, during the latter expedition, indicates that breeding took place here during 1998. Elephant seal counts from previous expeditions are summarised.     

Female competition and reproductive success in northern elephant seals

The probability of weaning a healthy pup increases with age in female northern elephant seals, Mirounga angustirostris. On Año Nuevo Island, California, weaning success among ‘prime’ females, those 6 years of age or older, was more than double that of ‘young’ females, those 3 to 5 years old. Prime females were better mothers than young females because of superior size, higher social dominance, and greater maternal experience; they were more likely to mate with high-ranking males and gave birth at an optimal time and place, circumstances that maximized the probability that their pups would survive, develop, and reproduce. The competitive advantage of prime-age mothers over younger ones was greatest when female and pup density was high. Young females improved their chances of reproducing successfully by emigrating from crowded harems and establishing new colonies.     

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.     

Vagrant elephant seal predation on Cape fur seal pups, Plettenberg Bay, South Africa

Vagrant southern elephant seals (Mirounga leonina) are occasionally sighted along the coast of South Africa and are known to feed primarily on fish and squid. Phocid seals are not known to predate on mammals making the events described here exceptional. This note describes the successful and failed attempts of a vagrant male southern elephant seal (M. leonina) to consume Cape fur seal (Arctocephalus pusillus pusillus) pups in Plettenberg Bay, South Africa. Observations were made by crew and passengers aboard a commercial whale-watching vessel during November 2012. This is the first account of elephant seals eating anything other than fish, squid and penguins and suggests considerable plasticity in prey choice dictated by environmental conditions.