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.     

Killing of Cape fur seal (Arctocephalus pusillus pusillus) pups by brown hyenas (Parahyaena brunnea) at mainland breeding colonies along the coastal Namib Desert

Brown hyenas (Parahyaena brunnea) scavenge and kill seal pups at mainland Cape fur seal (Arctocephalus pusillus pusillus) colonies. The prey encounter interval and interval between kills depended on seal density, and increased density resulted in an increase of the capture rate and increase in hunting efficiency from 14% in November to 47% in January. The time brown hyenas spent at the seal colony decreased with increasing seal density and increasing air temperatures. Nevertheless, they were regularly active during the day when less adult seals were present at the colony, which indicates that the attendance of adult seals might play a role in the choice of foraging time. Brown hyenas killed seal pups throughout the study period. The predation rate was independent of the availability of non-violent mortalities, but the absolute number of kills was positively density-dependent. Mass kill events were recorded throughout the study period and are therefore not unusual occurrences. The overabundance of easy and vulnerable prey may lead to an over stimulus situation that triggers killing independent of the consumption of the prey or the hunger state.     

Holocene changes in the ecology of northern fur seals: insights from stable isotopes and archaeofauna

The remains of northern fur seals (Callorhinus ursinus) are among the most abundant of pinniped elements recovered from mainland coastal archaeological sites in both California and Oregon. This is surprising as all contemporary northern fur seals breed exclusively on offshore islands, primarily at high latitudes, and the species is otherwise pelagic. The vulnerability of these animals to human predation suggests that either humans were foraging much further offshore than has been presumed or alternatively that the ecology of these animals has shifted during the late Holocene. We used isotopic and archaeofaunal analysis of the remains of pinnipeds from the middle to late Holocene of central and northern California to clarify the breeding and foraging behavior, and migration patterns of these ancient animals. The carbon and nitrogen isotope compositions of ancient northern fur seals reveal that these animals fed as far offshore as they do today, and that they remained at middle latitudes throughout the year. From an archaeological site at Moss Landing, California, we identified 16 skeletal elements from at least 12 very small northern fur seal pups. From another site near Mendocino, California, we identified the remains of at least 6 pups. We estimate the size and age of 5 of the young animals using sex-specific regressions of body length on the short dentary length derived from measurements of modern specimens. Our estimates indicate these ancient pups were substantially smaller, and therefore younger, than modern 3-month-old northern fur seal pups from similar latitudes and their nitrogen isotope compositions suggest they had not been weaned. As present-day northern fur seals do not leave their rookeries until they are at least 4 months old, we consider it highly unlikely that these ancient pups swam to these mainland locations from some distant island rookery. While there are numerous nearshore rocky outcrops along the Mendocino Coast, which may have supported small breeding colonies, the Moss Landing site is centered on a 40-km-long sandy beach, and is more than 120 km from what at the time were the nearest offshore islands. We conclude that northern fur seal adult females, subadults, and pups whose remains were recovered at the Moss Landing archaeological site must have been taken at a mainland rookery. Evidence that northern fur seals once bred on the mainland at this central California location suggests that the abundant remains of these animals at numerous other archaeological sites along the California coast also reflect the presence of nearby mainland rookeries. Based on the relative abundance of their remains in ancient human occupation sites and the widespread distribution of sites where their remains have been found, it appears that northern fur seals were once the predominant pinniped throughout a region where they now only rarely occur. Furthermore, their presence along the central and northern California coasts appears to have once severely limited the distribution of other pinnipeds, which are now common to the region.     

THERMAL FUNCTION OF PHOCID SEAL FUR

In phocid seals, blubber serves as the main thermal insulation instead of fur. The thermal function of fur, at least in adult phocid seals, has therefore been questioned. We measured the relative contribution of fur to the combined thermal resistance (insulation) offered by blubber, skin, and fur in newborn and adult harp ( Pagophilus groenlandicus ) and hooded ( Cystophora cristata ) seals, in air and water, to elucidate the role of fur as insulation in phocid seals. In air the fur contributed 90% of the combined thermal resistance of blubber, skin, and fur in newborn harp seal pups and 29% in adulrs, whereas in hooded seals the fur contributed 73% in newborn pups and 34% in adults. When submerged the thermal resistance of the fur was reduced by 84%-92%, and contributed 65% to the total insulation in newborn harp seal pups and 3% in adults, and 26% in newborn hooded seal pups and 5% in adults. We conclude that in air the fur of phocid seals makes an important contribution to the insulation of pups, and also contributes considerably to the insulation of adult animals. In water, even though the thermal resistance of the fur is dramatically reduced, the fur still contributes substantially to the insulation of pups, but its contribution in adults is negligible.     

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.     

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.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in relation to temporal variation in predation risk by white sharks (Carcharodon carcharias) around a seal rookery in False Bay,South Africa

The marked differences in predation risk posed by white sharks (Carcarodon carcarias) at island rookeries of Cape fur seals (Arctocephalus pusillus pusillus) offer a quasi‐experimental design within a natural system for exploring how prey adjust their behavior in response to temporal variation in predation risk. Here we compare movement of juvenile and adult Cape fur seals at a high risk (Seal Island) and low risk (Egg Island) rookery. We further compare juveniles and adults at Seal Island in low and high risk seasons and at low and high risk times of day within those seasons. Adult fur seals at Seal Island avoided traversing the zone of high white shark predation risk during the high risk period (0700–0959) in the season of high risk (winter), but not during the low risk season (summer). By contrast, adult fur seals at Egg Island showed no temporal discretion in either season. Unlike juvenile fur seals at Egg Island, juveniles at Seal Island adjusted their temporal movement patterns to more closely mimic adult seal movement patterns. This suggests that exposure to predators is the primary driver of temporal adjustments to movement by prey species commuting from a central place.     

Multiple mating strategies explain unexpected genetic mixing of New Zealand fur seals with two congenerics in a recently recolonized population

Human impacts on natural systems can cause local population extinctions, which may promote redistribution of taxa and secondary contact between divergent lineages. In mammalian populations that have mating systems shaped by polygyny and sexual selection, the potential for hybridization to ensue and persist depends on individual and demographic factors. At Macquarie Island, a recently formed fur seal population is comprised of both sexes of breeding Antarctic (Arctocephalus gazella) and subantarctic (A. tropicalis) fur seals, and an itinerant collection of male New Zealand fur seals (A. forsteri), presumed to be non-breeders due to their absence from principle breeding areas. The mating system of the three species is described as resource-defence polygyny: males defend beach territories containing breeding females for exclusive mating rights. A recent genetic study identified a high level of hybridization in the population (17-30%), unexpectedly involving all three species. This study examined the source of involvement in breeding by A. forsteri with respect to mating strategies operating in the population. Ninety-five (10%) pups born from 1992 to 2003 were genetically identified as New Zealand hybrids. Most resulted from reproduction within territories by New Zealand hybrids of both sexes, although some were conceived extra-territorially, indicating that males successfully utilize strategies other than territory holding to achieve paternities. Female reproductive status influenced mating partner and mating location, and females without pups were more likely to conceive extra-territorially and with A. forsteri males. This study illustrates an important consequence of low heterospecific discrimination in a sympatric population of long-lived mammals.     

Vagrant Antarctic pinnipeds at Gough Island

Opportunistic sightings of Antarctic fur seals Arctocephalus gazella and a leopard seal Hydrurga leptonyx at Gough Island (40°20′S, 09°54′W) in the South Atlantic Ocean represent the northernmost island records for vagrant Antarctic fur seals, and only the second for a vagrant leopard seal at Gough Island. Some ten different individual Antarctic fur seals were sighted on a single day, up to seven on a single beach. An overall total of 18 individuals were recorded over a 7-week period in October/November 2005, before the onset of the breeding (pupping) season of the resident population of Subantarctic fur seals Arctocephalus tropicalis. Most were immature, male and mostly in good condition (n = 11, 61%); one mature male and six putative females were also present. All had departed after 23 November, although only a fraction of the available beaches was searched.     

The contemporary condition and some demographic characteristics of the Steller sea lion (<Emphasis Type="Italic">Eumetopias jubatus</Emphasis>) reproductive group on Tyuleniy Island,Sea of Okhotsk

In 2010, the largest part of the Steller sea lion breeding community on Tyuleniy Island was located on the harem rookery of northern fur seals, which occupied the eastern beach, as well as on the western side of the island, which was free of fur seals. At the culmination of harem activity on June 29, 26.5% of the animals at the age of 1+ concentrated on the eastern beach and 41.1%, on the western beach in the daytime. However, 52.3% of the pups were born on the eastern beach and only 30.4% were born on the western beach. Pups were also present on the capes: 9.1% of the pups were observed on the northern cape and 8.2% on the southern cape, while the main population on these sites consisted of non-harem bulls, bachelors, and young animals. At the peak of harem activity, the number of females per one harem bull was 13.1 at sites 1 to 3 of the eastern beach and each of them, on average, had 1.05 pups; on sites 7–12 there were, respectively, 9.1 females and 1.42 pups per female, and on the western beach, 21.7 females and 0.64 pups. The resulting abundance of sea lions on Tyuleniy Island in 2010 exceeded 1500, which was almost ten times as many as their number in 1989. A total of about 100 bulls, 60 harem bulls, 1000 females, and 700 pups were recorded there. Half-bulls and young animals amounted to one-third of the entire population. Meanwhile the overall sex ratio at the culmination of harem activity was 11.5 females per one bull and 18.8 per one harem bull. About 75% of the females belonged to the parous group. The mortality rate among newborns reached 5.4%. No mortality was observed in adults. As many as 133 previously branded Steller sea lions were found and 109 of them (81.9%) were immigrants. Among immigrants, 29% were branded individuals of reproductive groups from the Kuril Islands, 54% were from the Iony Islands, 16% were from the Yamsky Islands, and about 1% were from Kamchatka. Four-year-old individuals predominated among the branded immigrants (23.8%). The oldest Steller sea lion (21 years of age) was one that was branded on the Srednego Islands in 1989. The rate of marked animal return from 175 pups that were branded on Tyuleniy Island the year before was 13.8%.     

Low reproductive success in territorial male Antarctic fur seals (Arctocephalus gazella) suggests the existence of alternative mating strategies

Microsatellites were used to conduct an analysis of paternity of Antarctic fur seals (Arctocephalus gazella) from Bird Island, South Georgia. At most, only 28% of pups at our study site could be assigned a father, even though the majority (approximately 90%) of candidate males within this colony were sampled. The behavioural and genetic evidence from this study suggests that a number of alternative mating strategies may exist within this fur seal population. Holding a land-based territory conferred an advantage to male reproductive success. However, this advantage was much smaller than expected from behavioural observations. At least 70% of fur seal pups born at our study site in a given year are not fathered by males who held a territory or were observed copulating with females in the previous year, implying that there exists a pool of males that seldom venture ashore at this site. To explain this discrepancy we suggest that female choice is an integral component of the Antarctic fur seal mating system and that aquatic mating may play a much larger role in the Antarctic fur seal than previously thought.     

Embryonic mortality and predation on egg capsules of the Port Jackson shark Heterodontus portusjacksoni (Meyer)

Egg capsules of the Port Jackson shark Heterodontus portusjacksoni were examined during underwater visual census surveys at two sites in New South Wales, Australia, during the 2002–2005 reproductive seasons (austral winter). Embryonic mortality was high (0·783–0·896 per annum) with the majority (99·2%) resulting from predation. The crested horn shark Heterodontus galeatus and common tent shell Astralium tentoriformis (Gastropoda, Turbinidae) were positively identified as egg predators. Gastropod predation was insignificant, accounting for only 2·0 and 2·1% of total mortality at Terrigal Haven and Dent Rock, respectively. Most capsules appeared to have been depredated by large predatory fishes with the eastern blue groper Achoerodus viridis (Labridae) and the black stingray Dasyatis thetidis (Dasyatidae) as possible candidate predators. The rate of embryonic mortality in H. portusjacksoni is higher than that reported for other oviparous elasmobranchs. This high level of mortality has significant consequences for the conservation and management of this species, especially when combined with an understanding of their low fecundity, late maturity and protracted reproductive life.     

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.     

DIET OF WEANED PUPS AND SEASONAL VARIATIONS IN BODY CONDITION OF JUVENILE BARENTS SEA HARP SEALS PHOCA GROENLANDICA

Data on blubber depth, body condition, and diet were collected from young of year and juvenile harp seals ( Phoca groenlandica ) in the Barents Sea in the periods February-April and June-October 1990–1997. Harp seal pups feed independently shortly after weaning. In the southern parts of the Barents Sea the diet of harp seal pups consisted mainly of euphausiids ("krill") Thysanoessa sp. and amphipods of the genus Parathemisto . The change in body condition of harp seal pups between weaning (mid-March) and June indicated that considerable amounts of blubber energy, deposited during suckling, were mobilized. This suggests that the early food intake of weaned pups was insufficient to meet the energy requirements. The pattern of low spring and earlysummer body condition, followed by a subsequent rapid improvement in condition during late summer and autumn appeared to occur not only among pups-of-the-year, but also in one- and two-year old harp seals.     

Grey seal, Halichoerus grypus, energetics: females invest more in male offspring

Grey seal females transfer large amounts of energy to their pups during the brief lactation period. The costs of lactation have been measured using weight changes of mother and pup pairs. Large females come ashore to give birth earlier in the season and lose weight more rapidly than smaller females. The sex ratio of Grey seal pups born is skewed towards males in the early part of the breeding season. Male pups are larger at birth and gain weight more rapidly than female pups, and their mothers show a correspondingly faster rate of weight loss than mothers of female pups. The energy costs of gestation and lactation to a Grey seal mother are 31 GJ for male pups and2–8 GJ for female pups. Males are therefore 10% more costly in energy terms to raise to weaning. Because, on average, large females arrive at breeding sites before smaller animals, biased results on weight changes would be obtained from methods which do not use repeated weighings. We suggest that the high efficiencies of lactation estimated for Harp seals compared with other phocid seals could be accounted for by such a bias.     

AGING LIVE ANTARCTIC FUR SEALS AND SOUTHERN ELEPHANT SEALS

This study describes a method for extracting post-canine or incisor teeth from live antarctic fur seals ( Arctocephalus gazella ) and southern elephant seals ( Mirounga leonina ) respectively and their use to determine age in a field situation. Dental elevators were used to loosen the teeth from the alveolus and periodental ligament. Most teeth were removed within l-2 min and a total of 214 and 81 teeth were collected by this method from antarctic fur seals and southern elephant seals respectively. No seal recaptured at intervals up to a year after a tooth was extracted showed signs of infection or distress related to removal of the tooth. Teeth were thin-sectioned for the purpose of aging. In both species cementum growth layer groups were a more satisfactory indicator of age than dentinal growth layer groups. Estimates of age from cementum growth layers were confirmed for Antarctic fur seals using seals which had been tagged as pups up to 16 yr before sampling.     

VOCAL INDIVIDUALITY IN MOTHER AND PUP SOUTH AMERICAN FUR SEALS, ARCTOCEPHALUS AUSTRALIS

South American fur seals breeding in Peru are subjected to levels of maternal aggression, and subsequent pup mortality, that are higher than has been reported for any other otariid species. For mothers and pups to maintain contact with each other, a mutual recognition system should exist to facilitate reunion and avoid misdirection of maternal effort. We recorded vocalizations of mothers and pups at Punta San Juan, Peru, during the 1994 and 1995 breeding seasons. Sixteen acoustic variables were measured from a total of 560 calls from 15 mothers and 13 pups. Multivariate analysis showed that calls were variable in several acoustic dimensions. While calls of both mothers and pups showed low variability within and high variability among individuals, mothers' calls were more individualistic. On average, discriminant-function analysis correctly assigned 60% of pup calls and 70% of mother calls to the individual that produced them. Characteristics of the fundamental frequency were most important for distinguishing among mothers, while pup calls, which typically contained less harmonic structure, could be differentiated by formant-like frequency ranges. Thus, calls of mother and pup South American fur seals appear to exhibit sufficient stereotypy to allow for recognition and discrimination among individuals.     

DISPERSAL and BYCATCH MORTALITY IN GRAY, HALICHOERUS GRYPUS, AND HARBOR, PHOCA VITULINA, SEALS TAGGED AT THE NORWEGIAN COAST

Between 1975 and 1998, 3,571 gray and 630 harbor seal pups were tagged along the Norwegian coast, and 259 (7%) gray and 80 (13%) harbor seal tags were returned. Incidental mortality, mainly in bottom-set nets, accounted for the majority of deaths (79% in gray and 48% in harbor seals, respectively). Seals were most vulnerable to incidental mortality in fishing gear during the first three months after birth, but high incidental mortality prevailed during the first 8–10 mo. Gray seals dispersed more widely (mean distance: 120 km) than harbor seals (mean distance: 69 km). Both species dispersed most widely during the two first months after tagging. The maximum distance moved was 739 km for gray and 463 km for harbor seals. Strong fidelity for their place of birth was observed in adult gray seals during breeding season. No significant difference in incidental mortality was detected between the areas of tagging. However, for 37 harbor seals tagged in a 724 km nature reserve no returns were reported.     

Low heritability and high phenotypic plasticity of salivary cortisol in response to environmental heterogeneity in a wild pinniped

Individuals are unique in how they interact with and respond to their environment. Correspondingly, unpredictable challenges or environmental stressors often produce an individualized response of the hypothalamic‐pituitary‐adrenal (HPA) axis and its downstream effector cortisol. We used a fully crossed, repeated measures design to investigate the factors shaping individual variation in baseline cortisol in Antarctic fur seal pups and their mothers. Saliva samples were collected from focal individuals at two breeding colonies, one with low and the other with high density, during two consecutive years of contrasting food availability. Mothers and pups were sampled concurrently at birth and shortly before weaning, while pups were additionally sampled every 20 days. We found that heritability was low for baseline cortisol, while within‐individual repeatability and among‐individual variability were high. A substantial proportion of the variation in baseline cortisol could be explained in pups and mothers by a combination of intrinsic and extrinsic factors including sex, weight, day, season, and colony of birth. Our findings provide detailed insights into the individualization of endocrine phenotypes and their genetic and environmental drivers in a wild pinniped. Furthermore, the strong associations between cortisol and life history traits that we report in fur seals could have important implications for understanding the population dynamics of species impacted by environmental change.     

Behavior of Cape fur seals (Arctocephalus pusillus pusillus) in response to spatial variation in white shark (Carcharodon carcharias) predation risk

Foraging and predation risk are often separated at rookeries of marine central place foragers, thus offering an opportunity to gain insight into how predator‐avoidance shapes the behavior of prey. Here we compare the behavior of Cape fur seals (Arctocephalus pusillus pusillus) at two island rookeries with and without white shark (Carcharodon carcharias) predations, and assess seal behavior in relation to marked spatiotemporal variation in risk at the high‐risk site (Seal Island, South Africa). Our results show that seal behavior at the two sites is comparatively similar in summer, when predation risk is low at both sites, but not in winter. Compared to seals at the “low‐risk” site, seals at Seal Island avoided deep‐water habitat around the island at high risk times and restricted their use of this habitat in favor of safe, shallow waters when engaging in social and thermoregulatory behaviors. Seals increased their frequency of jostling, porpoising, and diving when moving through the danger zone and seals in groups were safer than single individuals. Overall, our results suggest that seal behavior around the high‐risk site is strongly affected by predation risk, and show this rookery to be an excellent predator‐prey system at which to evaluate long‐standing ecological hypotheses.