Analysis of growth and stable isotopes in teeth of male Australian fur seals reveals interannual variability in prey resources

To detect and monitor long‐term ecosystem responses to environmental variability, managers must utilize reliable and quantitative techniques to predict future ecosystem responses. Canine teeth from 67 male Australian fur seals (aged 2–19 yr), collected at Seal Rocks, between 1967 and 1976, were measured for relative growth within the dentine growth layer groups (GLGs), as an index of body growth. Fluctuations in relative growth were apparent during 1956–1971, suggesting interannual variation in prey resources within Bass Strait. These were positively correlated with the Southern Oscillation Index and negatively with the Indian Ocean Subtropical Dipole, both on a 2 yr lag. The observed delay may reflect the time required for the nutrient cascade to filter through to the predominantly benthic prey of Australian fur seals. Stable isotope analysis (δ¹⁵N/δ¹³C) was also used to investigate whether fluctuations in growth were associated with differences in diet. Relative growth was found to be negatively correlated with δ¹⁵N, suggesting years of greater resource availability may be associated with individuals consuming proportionally more prey biomass of lower isotopic value. This study demonstrates that fluctuations in the dentine GLGs of male Australian fur seals are related to environmental parameters, suggesting variation in body growth is mediated by changes in prey resources.     

Characterization of Australian fur seal vocalizations during the breeding season

The vocal repertoire, structure, and behavioral context of airborne vocalizations produced by Australian fur seals (Arctocephalus pusillus doriferus) are described using recordings made at a breeding colony on Kanowna Island, Bass Strait, Australia. The study identified six different call types: three produced by males (bark, guttural threat, and submissive call); five produced by females (bark, guttural threat, submissive call, growl, and pup attraction call) and the female attraction call produced by pups and yearlings. Vocalizations were compared according to age and sex classes. The overall structure and function of the pup attraction and female attraction call produced by females, yearlings, and pups, was similar. However, while similar in their overall appearance, certain call types have a lower fundamental frequency when compared with other fur seals. In addition, the male bark call alters in rate of production according to the context used, where calls are slower when males are stationary and advertising their territorial status and faster when males are involved in confrontations with other males or actively herding females. Further research is required to investigate changes in environmental conditions and their effects on shaping the call structure and communication in Australian fur seals.     

Survival estimates for the Australian sea lion: Negative correlation of sea surface temperature with cohort survival to weaning

The Australian sea lion (Neophoca cinerea) population at Seal Bay Conservation Park, South Australia, is estimated to be declining at a rate of 1.14% per breeding season. To better understand the potential causes of this decline, survival rates were examined to 14 yr of age for eight cohorts marked as pups (aged 0.17 yr) between 1991 and 2002. Apparent yearly survival rates (Φ) varied by cohort for pups from marking to weaning at 1.5 yr (Φ= 0.30–0.67). Postweaning juvenile survival (1.5–3 yr) was 0.89 and survival from 3 to 14 yr was constant (Φ female:male = 0.96:0.89). Φ of pup cohorts was negatively correlated to local sea surface temperature where the sea lions forage (SST) and was especially low for cohort 7 in 2000 (0.30). It is possible that periods of unusually warm oceanographic conditions may be limiting primary production and inhibiting maternal provisioning to pups. Pup survival to weaning is relatively low compared to other otariid species, is likely to limit recruitment, and may be contributing to the decline in pup abundance observed in the colony.     

Temporal patterns of milk production in Antarctic fur seals (Arctocephalus gazella)

The timing of milk production in Antarctic fur seals was studied at Bird Island, South Georgia. Like all lactating otariid seals (Pinnipedia: Otariidae), Antarctic fur seal females alternate between short nursing periods ashore and regular foraging trips to sea. Females do not necessarily return to the colony with full mammae, which indicates that mammary volume capacity is unlikely to limit foraging trip duration. Upon arrival at the colony, milk fat (r²= 0.33, P < 0.04) and protein (r²= 0.60, P < 0.002) content were positively correlated to the time spent at sea. A similar trend was observed in the milk produced on land. The rate of milk energy production was much lower at sea (5.02 ± 0.05 MJ day^-1) than on land (23.66 ± 4.4 MJ^-1 day^-1). The rate of milk energy production during the foraging trip was negatively correlated to the time spent at sea (r²= 0.29, P < 0.05), whereas the rate of milk energy production on land was positively correlated (r²= 0.61, P < 0.001) to the duration of the preceding foraging trip. The total amount of milk energy delivered to the pup during each twoday nursing period was positively correlated (r²= 0.60, P < 0.002) to the duration of the previous foraging trip. The overall rate of milk energy delivery, however, was independent of foraging trip duration. This accords with previous observations that the growth rates of Antarctic fur seal pups are unaffected by maternal foraging trip duration patterns.     

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.     

The behavioural response of Australian fur seals to motor boat noise

Australian fur seals breed on thirteen islands located in the Bass Strait, Australia. Land access to these islands is restricted, minimising human presence but boat access is still permissible with limitations on approach distances. Thirty-two controlled noise exposure experiments were conducted on breeding Australian fur seals to determine their behavioural response to controlled in-air motor boat noise on Kanowna Island (39°10'S, 146°18'E). Our results show there were significant differences in the seals' behaviour at low (64-70 dB) versus high (75-85 dB) sound levels, with seals orientating themselves towards or physically moving away from the louder boat noise at three different sound levels. Furthermore, seals responded more aggressively with one another and were more alert when they heard louder boat noise. Australian fur seals demonstrated plasticity in their vocal responses to boat noise with calls being significantly different between the various sound intensities and barks tending to get faster as the boat noise got louder. These results suggest that Australian fur seals on Kanowna Island show behavioural disturbance to high level boat noise. Consequently, it is recommended that an appropriate level of received boat sound emissions at breeding fur seal colonies be below 74 dB and that these findings be taken into account when evaluating appropriate approach distances and speed limits for boats.     

AGE, GROWTH, AND REPRODUCTION IN NEW ZEALAND FUR SEALS

Growth and reproductive biology of New Zealand fur seals ( Arctocephalus forsteri ) were studied by examining 127 seals (57 females, 64 males) killed incidentally in fishing gear in New Zealand waters in 1996. Tooth sections were used to age the animals, and male and female reproductive organs were examined macroscopically and histologically. The maximum age observed was 22 yr for females and 12 yr for males. Males were significantly larger than females, but growth was similar up to 5 yr. Males reached sexual maturity between 5 and 9 yr of age, whereas females did so between 4 and 6 yr. The mean pregnancy rate in females was 0.69 (95% CI = 0.54–0.81).     

Factors affecting weaning weight in Antarctic fur seals Arctocephalus gazella at South Georgia

Summary Relationships between size, body condition, age and feeding-attendance patterns during pup rearing of female Antacrtic fur seals Arctocephalus gazella and their effects on the timing of birth and weaning, pup weight, growth and condition were studied at South Georgia in 1981–1982. Twenty-seven (6 male, 21 female) mother-pup pairs were followed from birth to weaning. The analysis of maternal effects was limited to female pups because of the small sample size of male weaners. High weaning weight was associated with those female pups whose mothers spent more time ashore attending their offspring. Weaning weight showed no relationship with perinatal duration, number of feeding trips to sea, days at sea or date of weaning. A further 63 mother-pup pairs were analysed for the effects of maternal body condition (weight/length), age and timing of birth on offspring body weight and condition. Pup weight and condition were weakly correlated with maternal age in female pups. Male pups born earlier in the season were heavier and in better condition. Maternal and offspring body weight and condition were unrelated. For the Antarctic fur seal population at South Georgia where the food supply was apparently not limiting in summer, maternal condition and foraging time were subordinate to maternal care on land (as expressed by attendance duration) in determining offspring weight at weaning.     

NEWS

ABSTRACT

Australian fur seals Arctocephalus pusillus doriferus are colonial breeding animals forming dense social groups during the breeding season. During this time, males establish and defend territories through physical conflicts, stereotyped posturing and vocalisations. While vocalisations are suggested to play an important role in male recognition systems, it has received little attention. Recordings of nine adult male Australian fur seals were made during the 2000 and 2001 breeding seasons at Kanowna Island (39° 10′S, 146° 18′ E), Bass Strait, Australia. The in-air bark vocalisations of territory-holding males were used to characterise the Bark Call and to determine whether males produce individually distinct calls, which could be used as a basis for vocal recognition. Seventeen frequency and temporal variables were measured from a total of 162 barks from nine individual males. The Bark Series was more reliably classified (83%) to the correct caller compared to the Bark Unit. This was assigned with less certainty (68%), although the classification was still relatively high. Findings from this study indicate that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition in male Australian fur seals.     

REDUCED POPULATION GROWTH OF GRAY SEALS AT SABLE ISLAND: EVIDENCE FROM PUP PRODUCTION AND AGE OF PRIMIPARITY

Pup production on Sable Island, Nova Scotia, has been increasing exponentially since the early 1960s and by 1997 Sable Island was the largest gray seal colony worldwide. Using an aerial photographic survey, as in previous years, we estimated pup production in January 2004 to determine if this exponential rate of increase had continued. A total of 33,268 pups was counted on the color positives. When corrected for the proportion pups missed on the imagery (1.106 for the 12th; 1.527 on the 13th), the proportion of pups that died prior to the survey (0.031), and the proportion of pups born before the survey (east colony 0.966, west colony 0.962), estimated total pup production was 41,500 with SE = 4,381. The 2004 estimate indicates that pup production on Sable Island has continued to increase, but suggests that the rate of increase ( r ) may have declined (0.070 compared to previous 0.128). Females from the 1998–2000 cohorts were about 16 times less likely to give birth for the first time at age 4 yr and more than twice as likely at age 6 yr compared to those in the mid-late 1980s. The new estimate of pup production and observed changes in age of primiparity provide the first indication of changes in vital rates of this population. However, additional estimates of pup production and vital rates are needed to confirm this conclusion and to investigate the underlying mechanisms.     

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.     

BODY TEMPERATURE IN THE NEWBORN SOUTHERN ELEPHANT SEAL, MIROUNGA LEONINA, AT MACQUARIE ISLAND

Newborn southern elephant seal pups were reported by Laws (1953) to be "to some extent poikilothermic at birth." Rectal temperatures of known age southern elephant seal pups were recorded during the 1985 pupping season at Macquarie Island. The mean pup rectal temperature was found to be 381°C ± 0.1°C SEM ( n = 131, range = 36.5°-39.1°C). Pups at two hours, six hours, and one day after birth had significantly higher rectal temperatures than pups two, three, or four days of age. Rectal temperatures of neonatal southern elephant seals were within the range observed for other pinnipeds, (but never as low as the 31°C previously observed for southern elephant seals at Signy Island in 1953). A significant though weak positive correlation was found between pup temperature and body weight. However, no correlation between pup temperature and age or any environmental factor was found. These observations demonstrated that southern elephant seal pups at Macquarie Island are homeothermic, rather than heterothermic from birth.     

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.     

Dynamic influence of maternal and pup traits on maternal care during lactation in an income breeder, the antarctic fur seal

Life-history theory predicts that selection will favor optimal levels of parental effort that balance benefits of current reproduction with costs to survival and future reproduction. The optimal level of effort depends on parental traits, offspring traits, and provisioning strategy. Additionally, how these factors influence effort may differ depending on the stage of reproduction. The relative importance of maternal and offspring traits on energy allocation to offspring was investigated in known-age Antarctic fur seals Arctocephalus gazella across four stages of reproduction, using birth mass and milk-consumption measurements. Maternal traits were important during three of the four stages investigated, with larger females giving birth to larger pups and investing more in pups during perinatal and molt stages. Pup mass influenced maternal effort during the premolt stage, and provisioning strategy influenced postnatal maternal effort at all stages. Energy provided to the offspring during an attendance visit was positively related to the duration of the foraging-trip/visit cycle; however, when investment was controlled for trip/visit cycle duration, the overall rate of energy transfer was similar across trip durations. In addition to strong effects of maternal mass, pup traits affected energy allocation, suggesting that pup demand is important in determining maternal care. These findings emphasize the importance of considering state variables in life-history studies and suggest that timing of measurements of effort in species with long provisioning periods may influence conclusions and our ability to make comparisons of reproductive effort among species.     

Pup weight and survival of northern fur seals Callorhinus ursinus

The relationship between northern fur seal pup weight and post-weaning survival is examined. Males that survived at least two years after weaning had been significantly heavier than the mean for their cohort as pups. Among a small sample of females no significant relationship was found. Cohort survival from weaning to age two was not significantly correlated with mean pup weight. Pup weight may influence post-weaning survival less than other factors such as gender and variability of the environment. To the degree that it determines survival, pup weight may reflect the reproductive success or fitness of parents. Weight differences between tagged and untagged pups explain, in part, differences observed in their survival.     

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.     

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%).     

A PROTRACTED FORAGING AND ATTENDANCE CYCLE IN FEMALE JUAN FERNÁNDEZ FUR SEALS

Previous studies of fur seals suggest that the attendance patterns and consequent temporal patterning of energy transfer from mother to pup follows a latitudinal cline. While data from subpolar, tropical, and some temperate latitude species support the postulated cline, data for the temperate latitude Juan Fernández fur seal do not. Maternal foraging trips and associated visits ashore were the longest of all otariids studied to date. The first foraging trip postpartum averaged 10.2 d ( n = 51 females, range 1–22.5), foraging trips combined averaged 12.3 d ( n = 100, range 1.0-25.0), and visits ashore averaged 5.3 d ( n = 91, range 0.3-15.8) over the three seasons of study. Only duration of lactation was intermediate between subpolar and tropical strategies as predicted. Dive records suggest that these females feed almost exclusively at night at depths of less than 10 m and at distances of more than 500 km offshore. The prey species of this fur seal, primarily myctophids and squid, migrate to the surface at night and are patchily distributed. Foraging trip length varied between years in conjunction with shifts in seasurface temperature and type of prey consumed. We suggest that distribution of prey, irrespective of latitude, dictates foraging patterns of fur seals and leads to the exceptionally long foraging trips and visits ashore observed in this species.     

Tooth growth in male Antarctic fur seals (Arctocephalus gazella) from South Georgia: an indicator of long-term growth history

Growth of upper canine teeth of male Antarctic fur seals ( Arctocephalus gazella ) which died of natural causes at Bird Island, South Georgia, was quantified from measurements of annual layers in longitudinal sections of teeth. Mean age at death was 7.69±0.07 years and this showed a small but significant increase through the period when samples were collected (1972/73–1988/89). There were significant correlations between morphometrics of teeth and those of seals, suggesting that tooth growth provided an indication of body growth. Tooth growth rate was lowest in seals which died early (age 4 years) and increased with age at death. Changes in the growth pattern of teeth suggested that fur seals which became sexually mature early also died early. Tooth growth layers deposited in each calendar year were compared with the expected layer depth based on a linear relationship between layer depth and age at which each layer was deposited. There was significant variation in the depth of tooth growth layers deposited in different years, suggesting that growth was greater in some years than others. No trends in cohort strengths were detected, but particularly poor years for growth were closely related to years in which reproductive performance was also observed to be low. Variations in growth from 1967/68 to 1987/88 were correlated significantly (P < 0.008) with the Southern Oscillation Index of climatic variation.     

BREEDING BIOLOGY OF SOUTHERN ELEPHANT SEALS IN PATAGONIA

Abstract: Elephant seals breed in Patagonia (Península Valdés, Argentina) from late August to early November, reaching peak numbers during the first week in October. Observations of this population over the past ten years yielded similar results. Eighty percent of the pups were born by 2 October. Most (96%) of 663 females marked during three breeding seasons gave birth to a pup. Females stayed on land a mean of 28 d, gave birth 6 d after arrival, nursed their pups for 22 d, and copulated a mean of 2.5 times 20 d after parturition and 2 d before departure. Copulations peaked during the third week in October. Males spent 57–80 d on land fasting and defending harems of up to 134 females (median 11–13 females, depending on year). Most (96%) marked females that gave birth ( n = 636) also weaned their pups successfully. Pup sex ratio was unity. Harems were smaller and breeding occurred about three weeks earlier in Patagonia than in other colonies. Thermal conditions, day length and food availability may explain clines in the timing of breeding events between populations, Other parameters of the breeding season for the expanding Patagonia colony are similar to those for declining southern elephant seal populations elsewhere.