POPULATION SIZE AND BREEDING SEASON OF THE ANTARCTIC FUR SEAL ARCTOCEPHALUS GAZELLA AT HEARD ISLAND—1987/88

Breeding colonies of the antarctic fur seal Arctocephalus gazella on Heard Island (53°10'S, 73°30'E) are situated on the sheltered northern and eastern coasts on flat vegetated terrain near streams and pools. Pupping in the 1987/88 summer began on 21 November, with 90% of births in 26 d. The median birth date was 11 December. Pup counts at Heard Island made in seven breeding seasons from 1962/63 to 1987/88 show an exponential rate of increase of 21%, which may be inflated due to undercounting in early years. The total of 248 births in 1987/88 represents an exponential increase of 37% since the previous year, but pups may have been undercounted then. Based on the number of pups born, the breeding population is estimated at 870–1,120. During the breeding season, the largest number of animals ashore was 835. Many non-breeding fur seals began hauling out from early January and 15,000 animals were estimated to be ashore by late February, a fat larger number than expected from the size of the breeding population. Both the breeding and non-breeding components of the population may be augmented by immigration. The source of immigrants may be undiscovered breeding colonies of this species in the northwestern sector of the Kerguelen Archipelago or the concentration at South Georgia. Further censuses are required at Heard Island to monitor the population growth.     

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.     

BODY WEIGHT and GROWTH OF JUVENILE MALE NORTHERN FUR SEALS, CALLORHINUS URSINUS

A mark-recapture study conducted in 1987–1992 provided weight measurements of juvenile male northern fur seals ( Callorhinus ursinus ) on St. Paul Island, Alaska, at ages ranging from approximately 1.5 mo to 5 yr. Males born in 1987 tended to weigh less at ages 3 and 4 yr than those born in other years. Weights of individuals at ages 2, 3, and 4 yr were significantly correlated with their weights as pups ( P < 0.05). Weights at ages 2 and 3, 3 and 4, and 4 and 5 yr were significantly correlated ( P < 0.001), although weight changes with age were highly variable. Data indicate that larger than average male pups born during 1987–1990 were more likely to survive, but this effect was less evident than among pups born during 1960–1965 when average pup weights were lower.     

How mothers find their pups in a colony of Antarctic fur seals

In Antarctic fur seals, Arctocephalus gazella, mothers must identify their own young among hundreds or even thousands of pups, if they are to invest in their own offspring and avoid misdirecting their parental care. When returning to their breeding colony from a foraging trip of several days at sea, mothers have to find and identify their young before suckling can occur. There appears to be little confusion about which pup belongs to a mother, and adoption is absent or rare. Using behavioral observations, we investigated the means by which female Antarctic fur seals identified their pups in a breeding colony of about 750 mother-pup pairs on Kerguelen Island. We evaluated the importance of vision, scent communication, vocalizations, and rendezvous locations as possible explanations of how mothers find their pups. Every pup that a mother examined, whether her own or not, exchanged naso-nasal inspection with her, suggesting a strong role for olfactory communication in individual recognition. Both mothers and pups called to each other, and mothers that searched for pups over a longer period gave more calls and encountered more pups. Thus, vocalizations may have been used to attract pups that might be offspring. Nursing usually occurred in the same place from the end of one maternal visit to the colony and the arrival at the beginning of the next visit, suggesting that nursing locations may serve as a meeting place, or rendezvous, for mothers and pups. These results suggest that finding pups is a two-stage process for females, in which pups for sampling are attracted by calls or examined at the previous nursing location, and then individual identification is made by olfactory cues.     

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.     

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     

Factors affecting early mortality in the South American fur seal (Arctocephalus australis) in Peru: density-related effects and predation

Early mortality in the South American fur seal ( Arctocephatus australis ) in Peru is considerably higher than that seen in any other population of fur seal; 31–49% in the first month as opposed to a maximum of 20% in other populations. In 1987 and 1988, pup mortality was found to be enhanced by density-related effects and predation on pups by the southern sea-lion ( Olaria byronia ). At a high density beach, where 60% of the fur seals bred, mortality correlated with the number of females ashore as did aggression. Female aggression was driven to high levels by a combination of high density and movement within the colony to thermoregulate. However, even at a low density, beach mortality remained high due to a high level of predation by male southern sea-lions (low density beach 5·0–8·3% of all pups; high density beach 0·2%). Predation rates were lower at the high density beach because male fur seals expelled most attacking sea-lions during the breeding season (low density beach 10·0% of attacking sea-lions expelled; high density beach 58·8%).     

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.     

Annual variation in the timing of reproduction in Antarctic fur seals, Arctocephalus gazella, at Bird Island, South Georgia

The arrival of Antarctic fur seals at a breeding beach on Bird Island, South Georgia, was studied over five consecutive breeding seasons, 1983 to 1987. Experienced bulls arrived first and established breeding territories on the beaches in anticipation of the arrival of the cows. Male arrival, which is less synchronous within years than female arrival, was significantly later in 1987 than in any other year. Female arrival, estimated by pup birth date where necessary, was highly synchronous; it usually started when 80% or more of potential territory sites were occupied by males. Cows arrived significantly later in 1984 and 1987 than in 1983, 1985 or 1986. The late arrival of both males and females in 1987 is attributed to unusually severe climatic conditions during the preceding winter. The late arrival and reduced fecundity of females in 1984 is attributed to markedly reduced food availability during the austral winter and summer of 1983. Males were not affected in 1984 because they could move away from the area of reduced food availability earlier than females and because they have a more varied diet. Factors influencing the winter distribution, the timing and pattern of arrival and the breeding of male and female Antarctic fur seals are discussed.     

Fish prey of Antarctic fur seals,Arctocephalus gazella,during the summer-autumn period at Laurie Island,South Orkney Islands

The fish component in the diet of Antarctic fur seals, Arctocephalus gazella, was investigated at Laurie Island, South Orkney Islands from mid January to April 1988. Fish otoliths occurred in 78.8% of faecal droppings. Most of the otoliths extracted from scats belonged to Myctophid fish (93.4%), mainly Electrona antarctica and Gymnoscopelus nicholsi. These two fish species thus constituted the bulk of the diet. A greater percentage of the otoliths from G. nicholsi (31.1%) in this study were considered suitable for measurement in comparison with 10.3% from scats at Heard Island in 1990 and 11.4% at Macquarie Island in 1988/89. This lesser degree of erosion would suggest that foraging areas of fur seals during this study were closer to land than during previous studies. During the period studied, the commercial fishery around the South Orkneys was not based on Myctophid fish so there was no direct competition for fish between the fur seals and fisheries.     

Early-life density-dependence effects on growth and survival in subantarctic fur seals

Understanding the regulation of natural populations has been a long-standing research program in ecology. Current knowledge on marine mammals and seabirds is biased toward the adult component of populations and lacking are studies investigating the juvenile component. Our goal was to estimate demographic parameters on the pre-weaning stage of a subantarctic fur seal (Arctocephalus tropicalis) population on Amsterdam Island, suspected to be regulated by density-dependence. The influence of abundance on growth parameters (length and weight) and survival was assessed over a study period spanning 16 years. We evidenced a negative trend in population growth rate when density increased. Density-dependence models were favored for pup body size and mass growth. Abundance had a clear influence on body length at high population-density, pups grew slower and were smaller at weaning than pups born in years with low population density. Abundance partly explained pup body mass variation and a weak effect was detected on pre-weaning survival. The causal mechanisms may be increased competition for food resources between breeding females, leading to a reduction of maternal input to their pups. Our results suggested that pup favored survival over growth and the development of their diving abilities in order to withstand the extreme fasting periods that are characteristic of this fur seal population. This analysis provides significant insight of density-dependent processes on early-life demographic parameters of a long lived and top-predator species, and more specifically on the pre-weaning stage with important consequences for our understanding of individual long-term fitness and population dynamics.     

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.     

Inter-island variation in the diet of female northern fur seals (Callorhinus ursinus) in the Bering Sea

The diet of adult female northern fur seals ( Callorhinus ursinus ) is examined through the analysis of faecal material collected during the summer breeding season at three breeding locations in the Bering Sea: St. Paul Island (1988, 1990) and St. George Island (1988, 1990) of the Pribilof Islands Group (USA), and Medny Island (1990) of the Commander Islands Group (Russia). Prey consumption varies annually and accordingly with the physical and biological environment surrounding each island. Juvenile walleye pollock ( Theragra chalcogramma ) is the most common prey of northern fur seals from St. Paul Island; the island is surrounded by a broad neritic environment with widely separated frontal zones and is the greatest distance from the continental shelf-edge. Gonatid squid ( Gonatopsis borealis/Berryteuthis magister and Gonatus madokail Gonatus middendorffi ) were the most common prey of northern fur seals from Medny Island; the island is surrounded by a compressed neritic environment and is adjacent to the continental shelf-edge and the oceanic marine environment. A combination of walleye pollock and gonatid squid is consumed by northern fur seals from St. George Island; the island has a surrounding oceanographic environment intermediate between the other two islands.
Variability in predation on walleye pollock is consistent with fishery information concerning the relative abundance and availability of walleye pollock around St. George and St. Paul Islands. The abundance and availability of these prey resources during the summer breeding season are key factors which influence the health and growth of the northern fur seal populations in the Bering Sea.     

Ménage à trois on Macquarie Island: hybridization among three species of fur seal (Arctocephalus spp.) following historical population extinction

Human-induced changes to natural systems can cause major disturbances to fundamental ecological and population processes and result in local extinctions and secondary contacts between formerly isolated species. Extensive fur seal harvesting during the nineteenth century on Macquarie Island (subantarctic) resulted in extinction of the original population. Recolonization by three species has been slow and complex, characterized by the establishment of breeding groups of Antarctic and subantarctic fur seals (Arctocephalus gazella and Arctocephalus tropicalis) and presumed nonbreeding (itinerant) male New Zealand fur seals (Arctocephalus forsteri). One thousand and seven pups from eight annual cohorts (1992-2003) were analysed using mitochondrial control region data (RFLP) and 10 microsatellite loci to estimate species composition and hybridization. Antarctic fur seals predominated, but hybridization occurred between all three species (17-30% of all pups). Involvement of New Zealand fur seals was unexpected as females are absent and males are not observed to hold territories during the breeding season. The proportion of hybrids in the population has fallen over time, apparently owing to substantial influxes of pure Antarctic and subantarctic individuals and non-random mating. Over 50% of New Zealand hybrids and 43% of Antarctic-subantarctic hybrids were not F(1), which indicates some degree of hybrid reproductive success, and this may be underestimated: simulations showed that hybrids become virtually undetectable by the third generation of backcrossing. While human impacts seem to have driven novel hybridization in this population, the present 'time slices' analysis suggests some biological resistance to complete homogenization.     

Trend changes in sympatric Subantarctic and Antarctic fur seal pup populations at Marion Island,Southern Ocean

Recent pup population estimates of sympatric Subantarctic (Arctocephalus tropicalis) and Antarctic fur seals (A. gazella) at Marion Island are presented. Published pup population estimates of A. tropicalis (1995 and 2004) with an unpublished total island count in 2013, and annual counts on subsets of rookeries (2007–2015) were analyzed using a hierarchical Bayesian model. The pup population declined by 46% (95% credible interval CI: 43%–48%) between 2004 (mean = 15,260, CI: 14,447–16,169 pups) and 2013 (mean = 8,312, CI: 7,983–8,697), mirrored by a 58%–60% decline at rookeries counted annually (2007–2015). Population decline was highest at high‐density west and north coast rookeries, despite negligible change in female attendance patterns, pup mortality or median pupping date over the previous 25 yr. A better understanding of foraging behavior and its effects on reproductive success and survival in this A. tropicalis population is needed before we can attribute population decline to any external factors. In contrast, total island counts of A. gazella pups in 2007, 2010, and 2013, suggest that this population is still increasing although the annual intrinsic rate of population growth decreased from 17.0% (1995–2004, 744 pups) to 4.0% (2010–2013, 1,553 pups). The slowed growth of A. gazella is likely the result of saturation at the main rookery.     

Pup growth in the fur seals Arctocephalus tropicalis and A. gazella on Marion Island

Subantarctic fur seal Arctocephalus tropicalis and Antarctic fur seal A. gazella pups at Marion Island were weighed frequently until weaning, which occurs earlier in A. gazella (112 days) than in A. tropicalis (± 300 days). The mean birth weight of both species was the same (4·2 kg) and males grew faster than females. Arctocephalus gazella growth was linear to weaning and faster than A. tropicalis growth, which was linear to 120 days of age, slowing until 203 days of age, and thereafter losing weight but recovering perceptibly prior to weaning. Tagging had no apparent effect on pup growth. Faster growth in males than in females is part of the differential growth patterns which lead to adult sexual dimorphism. Arctocephalus gazella pup growth at Marion Island is faster than at South Georgia, indicating that conditions at this, their most northerly breeding locality, are not limiting for pup growth. The decrease in A. tropicalis pup body weight in July/August may result from either a scarcity of food in winter or weaning having been initiated. The two groups (polar and non-polar) of fur seals exhibit two strategies, which include differences in pup growth and suckling period; the polar species have a short suckling period and rapid pup growth and are predominantly pelagic over winter, while the more temperate species have a longer suckling period and slower pup growth and are less pelagic over winter.     

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.     

Mathematical modeling of the variation in the survival of female northern fur seals, <Emphasis Type="Italic">Callorhinus ursinus</Emphasis> (Linnaeus, 1758), on Tyuleniy Island

The dynamics of the survival and number of female northern fur seals, Callorhinus ursinus, on Tyuleniy Island have been studied. The data on the age structure and physiological condition of females (obtained during observations in 1958–1988) were used to estimate the model parameters. The adequacy of the model estimates of female number was evaluated using the data on the number of newborn pups at the rookery in 1958–2013. It has been found that a reduction in the survival rate of individuals of younger age groups, a decrease in the proportion of females, as well as aging of the population occurred in the 1958–1988 period. The use of numerical simulation methods shows that the rate of survival of females among individuals under 3 years of age increased substantially after 1988, indicating a positive trend in the dynamics of their number.     

PUP PRODUCTION AND POPULATION TRENDS OF THE AUSTRALIAN FUR SEAL (ARCTOCEPHALUS PUSILLUS DORIFERUS)

We estimated the number of live Australian fur seal pups using capture-markresights, direct ground counts, or aerial photography at all breeding sites following the pupping season of November-December 2002. Pups were recorded at 17 locations; nine previously known colony sites, one newly recognized colony and seven haul-out sites where pups are occasionally born. In order of size, the colonies were Lady Julia Percy Island (5,899 pups), Seal Rocks (4,882), The Skerries (2,486), Judgment Rocks (2,427), Kanowna Island (2,301), Moriarty Rocks (1,007), Reid Rocks (384), West Moncoeur Island (257), and Tenth Island (124). The newly recognized site was Rag Island, in the Cliffy Group, where we recorded 30 pups. We also recorded pups at the following haul-out sites: Cape Bridge-water (7 pups), Bull Rock (7), Wright Rock (5), Twin Islet (1), The Friars (1), He des Phoques (1), and Montague Island (1). In total, we estimate there were 19,819 (SE = 163) live pups at the time of the counts. We discuss trends in pup numbers and derive current population estimates for the Australian fur seal.     

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.