Decadal changes in habitat characteristics influence population trajectories of southern elephant seals

Understanding divergent biological responses to climate change is important for predicting ecosystem level consequences. We use species habitat models to predict the winter foraging habitats of female southern elephant seals and investigate how changes in environmental variables within these habitats may be related to observed decreases in the Macquarie Island population. There were three main groups of seals that specialized in different ocean realms (the sub‐Antarctic, the Ross Sea and the Victoria Land Coast). The physical and climate attributes (e.g. wind strength, sea surface height, ocean current strength) varied amongst the realms and also displayed different temporal trends over the last two to four decades. Most notably, sea ice extent increased on average in the Victoria Land realm while it decreased overall in the Ross Sea realm. Using a species distribution model relating mean residence times (time spent in each 50 × 50 km grid cell) to 9 climate and physical co‐variates, we developed spatial predictions of residence time to identify the core regions used by the seals across the Southern Ocean from 120°E to 120°W. Population size at Macquarie Island was negatively correlated with ice concentration within the core habitat of seals using the Victoria Land Coast and the Ross Sea. Sea ice extent and concentration is predicted to continue to change in the Southern Ocean, having unknown consequences for the biota of the region. The proportion of Macquarie Island females (40%) utilizing the relatively stable sub‐Antarctic region, may buffer this population against longer‐term regional changes in habitat quality, but the Macquarie Island population has persistently decreased (?1.45% per annum) over seven decades indicating that environmental changes in the Antarctic are acting on the remaining 60% of the population to impose a long‐term population decline in a top Southern Ocean predator.     

Rapid Response of a Marine Mammal Species to Holocene Climate and Habitat Change

Environmental change drives demographic and evolutionary processes that determine diversity within and among species. Tracking these processes during periods of change reveals mechanisms for the establishment of populations and provides predictive data on response to potential future impacts, including those caused by anthropogenic climate change. Here we show how a highly mobile marine species responded to the gain and loss of new breeding habitat. Southern elephant seal, Mirounga leonina, remains were found along the Victoria Land Coast (VLC) in the Ross Sea, Antarctica, 2,500 km from the nearest extant breeding site on Macquarie Island (MQ). This habitat was released after retreat of the grounded ice sheet in the Ross Sea Embayment 7,500–8,000 cal YBP, and is within the range of modern foraging excursions from the MQ colony. Using ancient mtDNA and coalescent models, we tracked the population dynamics of the now extinct VLC colony and the connectivity between this and extant breeding sites. We found a clear expansion signal in the VLC population ~8,000 YBP, followed by directional migration away from VLC and the loss of diversity at ~1,000 YBP, when sea ice is thought to have expanded. Our data suggest that VLC seals came initially from MQ and that some returned there once the VLC habitat was lost, ~7,000 years later. We track the founder-extinction dynamics of a population from inception to extinction in the context of Holocene climate change and present evidence that an unexpectedly diverse, differentiated breeding population was founded from a distant source population soon after habitat became available.     

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.     

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.     

Dinucleotide microsatellite markers from the Antarctic seals and their use in other Pinnipeds

Twenty‐four microsatellite loci were isolated from three species of Antarctic seals (Subfamily Monachinae, Tribe Lobodontini). Eleven loci were cloned from Weddell seal, Leptonychotes weddellii, seven from leopard seal, Hydrurga leptonyx, and six from crabeater seal, Lobodon carcinophagus. Variability was assessed in Weddell seals collected in McMurdo Sound, leopard seals from Bird Island, South Georgia, and crabeater seals sampled in the eastern Ross Sea. All loci were variable in the three species used for cloning and 22 of these loci amplified variable products in the Ross seal, Ommatophoca rossii. Cross‐species amplification was largely successful, with an average of 19 loci amplifying products in other phocids.     

Limited use of sea ice by the Ross seal (<Emphasis Type="Italic">Ommatophoca rossii</Emphasis>), in Amundsen Sea,Antarctica, using telemetry and remote sensing data

To understand the use and importance of the Antarctic sea ice to the Ross seal (Ommatophoca rossii), four adult females were tagged with Argos satellite transmitters in the Amundsen Sea, Antarctica. The Ross seal is the least studied of the Antarctic seal species and nothing was previously known about their behaviour in the Amundsen Sea. During almost 1 year, their movements, haul out behaviour and time spent at different temperatures were logged. By comparing their movements with daily ice maps, distances to the ice edge were calculated, and seals dependence on sea ice for resting, breeding and moulting was analysed. The tagged seals spent on average 70.8 % (range 66.8–77.8 %) of their time in the water and hauled out mainly during the moult in December–January, and in late October–mid-November during breeding. During the pelagic period, they were on average 837.5 km (range 587–1,282 km) from the ice edge indicating a fully pelagic life during several months. Their pelagic behaviour suggests that Ross seals, although being an ice obligate species, may adapt comparatively easy to climate change involving ice melting and recession and thereby potentially being less sensitive to the reduction of sea ice than other Antarctic seal species. Although nothing is known about their mating behaviour, they appear to be relatively stationary during moulting and breeding, hence requiring a small ice surface. Although previous studies in other parts of Antarctica have found similar results, still many questions remain about this peculiar species.     

Long-term monitoring of Antarctic pinnipeds in Admiralty Bay (South Shetlands, Antarctica)

Year-round monitoring of five Antarctic pinnipeds was conducted in Admiralty Bay from 1988 up to 2000. Two breeding species: southern elephant sealsMirounga leonina (Linnaeus, 1758) and Weddell sealsLeptonychotes weddellii (Lesson, 1826), were present throughout the year. Three other species: crabeater seals Lobodon carcinophagus (Hobron and Jacquinot, 1842), leopard sealsHydrurga leptonyx (Blainville, 1820), and Antarctic fur sealsArctocephalus gazella (Peters, 1875) visited the area only for short periods. During this study, the abundance of elephant seals was stable, whereas those of Weddell and crabeater seals declined. Leopard seals numbers fluctuated irregularly. We detected a possible immigration from South Georgia: of a stable magnitude for elephant seals, and of variable magnitude, depending on food accessibility, for Antarctic fur seals. We found a strong recurrence of the spatial distributions of elephant, Weddell, and Antarctic fur seals in the 13 oases on the shore of Admiralty Bay. Annual distribution patterns were characteristic for each species. The innermost beaches were used predominantly by the animals during their annual fasts: the breeding and the moulting seasons.     

基于基因组SNPs的南极恩克斯堡岛阿德利企鹅繁殖种群的遗传结构

我国提议在南极恩克斯堡岛新站址北侧3 km的阿德利企鹅(Pygoscelis adeliae)聚集繁殖地建立南极特别保护区, 对保护区边界的划分, 各国尚存争议, 尤其是对南湾(South Bay)的繁殖小种群是否具有遗传独特性, 是否应将其纳入保护区是重点关注的问题。本研究采集了恩克斯堡岛海景湾(Seaview Bay)和南湾的阿德利企鹅样品, 通过全基因组重测序和种群基因组学方法, 分析了恩克斯堡岛不同区域的种群遗传结构。发现恩克斯堡岛海景湾与南湾阿德利企鹅没有显著的遗传分化, 南湾阿德利企鹅不是独特的小种群; 海景湾高海拔区域个体与低海拔区域个体之间也没有显著的遗传差异, 推测该区域阿德利企鹅繁殖群体的分布格局与冰川堆积形成的阶地不具有显著相关性。本工作为恩克斯堡岛保护区和罗斯海新站建设提供了重要科技支撑。     

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.     

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 and breeding season chronology of Heard Island fur seals

Fur seals were eliminated by sealers at Heard Island soon after its discovery in the 1850s. The first recorded breeding of Antarctic fur seals (Arctocephalus gazella) since sealing was reported in early 1963 (two pups). The most recent survey of the Heard Island fur-seal population was undertaken between November 2000 and March 2001, when 1,012 Antarctic fur-seal pups were born. This represents a fourfold increase since the last complete census in 1987/1988 (13 years), when 248 births were recorded. Pup estimates and counts available for eight breeding seasons since 1962/1963 suggest the population has been increasing at between 12 and 20% per year. Based on pup production, the breeding population is estimated to number approximately 4,100 seals. The number of fur seals on Heard Island peaked in late February/early March at 29,256 indicating that, in addition to the breeding population, a significant number of seals born elsewhere haul out on the island. Most of these are moulting sub-adult and adult males. As in 1987/1988, only one subantarctic fur-seal pup (A. tropicalis) was observed, suggesting this species is not colonising the island, as has been speculated.     

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.     

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.     

An apparent population decrease,or change in distribution,of Weddell seals along the Victoria Land coast

Ground counts during 1959–1968 compared with counts using high resolution (0.6 m²) satellite imagery during 2008–2012 indicated many fewer Weddell seals (Leptonychotes weddellii) at two major molting areas in the western Ross Sea: Edisto Inlet‐Moubray Bay, northern Victoria Land, and McMurdo Sound, southern Victoria Land. Breeding seals have largely disappeared from Edisto‐Moubray, though the breeding population in McMurdo Sound appears to have recovered from harvest in the 1960s. The timing of decline, or perhaps spreading (lower numbers of seals in more places), is unknown but appears unrelated to changes in sea ice conditions. We analyzed both historic and satellite‐derived ice data confirming a large expansion of pack ice mostly offshore of the Ross Sea, and not over the continental shelf (main Weddell seal habitat), and a thinning of fast ice along Victoria Land (conceivably beneficial to seals). Timing of fast ice presence and extent in coves and bays along Victoria Land, remains the same. The reduction in numbers is consistent with an altered food web, the reasons for which are complex. In the context of a recent industrial fishery targeting a seal prey species, a large‐scale seal monitoring program is required to increase understanding of seal population changes.     

Minimum population size estimates demonstrate an increase in southern elephant seals (Mirounga leonina) on Livingston Island, maritime Antarctica

Southern elephant seals (Mirounga leonina) are apex predators of marine Antarctic food webs, and variations in their populations have been linked to environmental changes. Consequently, measuring and reporting the status of elephant seal populations provide insights into the environmental status of Antarctica. Here, we present new information on the size of the elephant seal subpopulation on Byers Peninsula (Livingston Island, South Shetland Islands, maritime Antarctica). Based on a total count of 1,510 pups, we estimated a total subpopulation size of 5,530 individuals by using a conversion factor of 3.5. This represents an increase of 150 % since the subpopulation was first counted 30 years ago. Based on this finding, we hypothesize that the overall South Georgia stock, to which the subpopulation we estimated on Byers Peninsula belongs, could be increasing instead of remaining stable as previously thought.     

Long-term variability in the abundance of Antarctic fur seals Arctocephalus gazella at Signy Island, South Orkneys

The number of Antarctic fur seals Arctocephalus gazella hauled out at Signy Island in the South Orkneys was monitored annually between 1977 and 2008. Over the study period seal abundance showed a tenfold increase, from a minimum of 1,643 seals in 1978 to a maximum of 21,303 in 1994. The majority of individuals observed were young adult males, likely to be migrants from South Georgia, with small numbers of female seals and only 65 pups recorded during the survey period. Variability in counts showed a similar pattern to Laurie Island, also in the South Orkneys archipelago, suggesting a similar annual immigration of seals to these two islands. The date of first seal arrival was correlated with the date of fast-ice breakout at Factory Cove, Signy Island, and years in which break out was exceptionally late (>21 December) corresponded with years of reduced seal abundance. While the presence of fast-ice during the early breeding season may currently inhibit the establishment of a major breeding population of fur seals at Signy Island, it is important that routine monitoring should continue, particularly in the light of current patterns of climate warming in the Antarctic.     

Vagrant Antarctic fur seals at Gough Island in 2009

The Antarctic fur seal Arctocephalus gazella at Gough Island (40°20′S, 09°54′W) in the South Atlantic Ocean, first seen in October/November 2005, was recorded again in September–October 2009. Up to three different individual Antarctic fur seals were sighted on a single day, on a particular beach. A total of seven different individuals were recorded over a 3-week period, well before the onset of the breeding (pupping) season of the resident population of Subantarctic fur seals A. tropicalis. Positively identified individuals were all male, mostly subadult and lean. Only a fraction (~20%) of the available beaches was searched, and it is unknown if the Antarctic fur seals were still present at Gough Island during the austral summer breeding season of southern fur seals.     

COLONIZATION HISTORY OF THE BALTIC HARBOR SEALS: INTEGRATING ARCHAEOLOGICAL, BEHAVIORAL, AND GENETIC DATA

Detailed knowledge about the history of colonization, population dynamics and behavior greatly enhance evaluation of genetic models of population units and migration rates in spatially structured populations. Here, the genetic uniqueness of harbor seals ( Phoca vitulinia ) in the eastern Baltic is evaluated in the light of new information on the distribution and abundance of Baltic and eastern North Sea populations during the last 11,000 yr, recent hunting statistics, and population counts. Archaeological records reveal that the Baltic population of harbor seals was founded about 8,000 yr ago. Adjacent populations in the North Sea areas were either small, or went extinct, and became significant only during the last 300 yr. This information generates the hypothesis that the Baltic population has been isolated during the last 8,000 yr, despite the lack of geographical barriers. We show that stochastic effects, isolation, and a documented recent population bottleneck can account for the low observed genetic variation in Baltic harbor seals.     

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.     

Climate Change Winners: Receding Ice Fields Facilitate Colony Expansion and Altered Dynamics in an Adélie Penguin Metapopulation

There will be winners and losers as climate change alters the habitats of polar organisms. For an Adélie penguin (Pygoscelis adeliae) colony on Beaufort Island (Beaufort), part of a cluster of colonies in the southern Ross Sea, we report a recent population increase in response to increased nesting habitat as glaciers have receded. Emigration rates of birds banded as chicks on Beaufort to colonies on nearby Ross Island decreased after 2005 as available habitat on Beaufort increased, leading to altered dynamics of the metapopulation. Using aerial photography beginning in 1958 and modern satellite imagery, we measured change in area of available nesting habitat and population size of the Beaufort colony. Population size varied with available habitat, and both increased rapidly since the 1990s. In accord with glacial retreat, summer temperatures at nearby McMurdo Station increased by ∼0.50°C per decade since the mid-1980s. Although the Ross Sea is likely to be the last ocean with an intact ecosystem, the recent retreat of ice fields at Beaufort that resulted in increased breeding habitat exemplifies a process that has been underway in the Ross Sea during the entire Holocene. Furthermore, our results are in line with predictions that major ice shelves and glaciers will retreat rapidly elsewhere in the Antarctic, potentially leading to increased breeding habitat for Adélie penguins. Results further indicated that satellite imagery may be used to estimate large changes in Adélie penguin populations, facilitating our understanding of metapopulation dynamics and environmental factors that influence regional populations.