Acoustic features involved in the neighbour-stranger vocal recognition process in male Australian fur seals

Many territorial species have the ability to recognise neighbours from stranger individuals. If the neighbouring individual is assumed to pose less of a threat, the territorial individual responds less and avoids unnecessary confrontations with familiar individuals at established boundaries, thus avoiding the costly energy expenditure associated with fighting. Territorial male Australian fur seals respond more to strangers than to neighbouring males. The present study evaluated which acoustic features were important in the neighbour-stranger recognition process in male Australian fur seals. The results reveal that there was an increase in response strength or intensity from males when they heard more bark units, indicating the importance of repetition to detect a caller. However, lengthening and shortening the inter-unit spaces, (i.e. changing the rhythm of the call) did not appear to significantly affect an animal's response. In addition, the whole frequency spectrum was considered important to recognition with results suggesting that they may vary in their importance. A call containing the dominant and surrounding harmonics was considered important to a male's ability to recognise its neighbour. Furthermore, recognition occurs even with a partial bark, but males need to hear between 25 and 75% of each bark unit from neighbouring seals. Our study highlights which acoustic features induce stronger or weaker responses from territorial males, decoding the important features in neighbour-stranger recognition.     

Ecological implications of body composition and thermal capabilities in young antarctic fur seals (Arctocephalus gazella)

In comparison with other homeotherms, young recently weaned marine mammals in high latitudes face exceptional energetic demands when foraging and thermoregulating. Lipids are an important source of energy and a major component of insulation that allows them to meet these demands. To examine the role of lipid stores in a high-latitude pinniped, we measured the body composition and thermoregulatory capabilities of Antarctic fur seal (Arctocephalus gazella) pups and yearlings by using flow-through respirometry and hydrogen isotope dilution. From these data, we constructed a model to examine the importance of postweaning fasting capability in free-ranging young fur seals. Resting metabolic rates were different for pups and yearlings measured in 0.6 degrees C water, 10.3 degrees C water, and ambient air; however, mass and percent lipid as covariates accounted for the different metabolic responses in pups and yearlings for all treatments. The estimated lower critical temperature for combined pups and yearlings was 14.4 degrees C, 10 degrees -15 degrees C above water temperatures normally experienced by Antarctic fur seals. Modeling predicted that a weaned fur seal pup would survive at sea from 9.8 to 36.2 d before succumbing to starvation. The most likely maximum travel distance within this time constraint suggests that food resources close to the natal rookery are important to first-year survival for this species.     

INDIVIDUAL VARIATION OF THE FEMALE ATTRACTION CALL PRODUCED BY AUSTRALIAN FUR SEAL PUPS THROUGHOUT THE MATERNAL DEPENDENCE PERIOD

ABSTRACT

Australian fur seals are colonial breeding animals that give birth on crowded rookeries. Females leave their pups unattended for extended periods while they forage at sea. On return to the colony, a mother must relocate her pup amongst the hundreds of other pups. Vocal recognition is thought to play a vital role in maintaining the mother-pup bond. In the present study, four sets of 7 different pups were recorded once each at different times throughout the maternal dependence period. The Female Attraction Call was used to determine whether Australian fur seal pups produce individually distinct calls which could be used as a basis for vocal recognition. Results from three different analyses (PIC, DFA and CART) indicated that the fundamental frequency, number of parts per call, duration, quavering and peak frequency changes at the start, mid-point and end of the call (i.e., along PEAK Fl) were important to recognition. In 75% of cases using DFA, the Female Attraction Call was classified to the correct caller, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by females.     

Distribution of post-weaning Antarctic fur seal Arctocephalus gazella pups at South Georgia

Potentially some of the biggest gaps in our knowledge about the ecology of Antarctic fur seals (Arctocephalus gazella) relate to juvenile animals. We investigated the at-sea distribution of five male and five female fur seal pups post-weaning. The study was carried out at Bird Island, South Georgia during two successive winters using satellite-linked platform transmitter terminals (PTTs). Our results are analysed in relation to pup sex and the physical environment and productivity of those 2 years, as well as in the context of our present knowledge of where post-breeding females and males forage. The available physical and biological data during both of the winters of this study suggest that both years were not unusual. We report marked differences between the sexes with male pups foraging significantly further away from land and their birth site than do females. The pups foraged in areas to the East of Bird Island seldom reported as foraging areas for the adult population. Also as winter progressed they showed a more oceanic distribution leaving the continental shelf, possibly to exploit a different prey source that was more readily available in the upper water column.     

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.     

Field metabolic rate of lactating female Galápagos fur seals (Arctocephalus galapagoensis): the influence of offspring age and environment.

Galápagos fur seals, Arctocephalus galapagoensis, live in a seasonal environment which varies strongly in productivity from year to year. We measured how the field metabolic rates (FMR) of lactating females varied with season, pup age and year. Energy expenditure was measured using doubly labeled water (DLW) during the cold seasons of 1984 and 1985 in 9 mothers of 1-3-month-old pups and 5 mothers of yearlings, and during the 1986 warm season in 8 mothers of 6-month-old pups. Young pups gained 0.84% mass/day during the cold season, but larger pups during the warm season lost 1.25% mass/day. During the warm season, females had lower relative total body water than during the cold season suggesting higher fat content during the warm, less productive season, but the effect was even more marked when comparing different years of the study: fat content was high in 1984 and 1986 and low in 1985. The FMR of mothers varied from 134 to 167 W but did not show significant differences between any of the pup age-groups. Among the years of the study, FMR showed only a trend towards low energy expenditure of mothers of young pups in 1984. The mean FMR was lower than for other otariids. Mothers may limit energy expenditure independent of pup age and season to minimize their own risk of starvation in an environment of comparatively low productivity, varying unpredictably due to frequent El Ni?o events.     

Observations on the infectivity of parasitic third-stage larvae of Uncinaria lucasi Stiles 1901 (Nematoda: Ancylostomatidae) of Northern fur seals, Callorhinus ursinus Linn., on St. Paul Island, Alaska.

Twelve fur seal pups, which had not nursed their mothers, were used in an infectivity experiment. Pups were exposed to parasitic 3rd-stage larvae of Uncinaria lucasi from belly tissues of fur seal bulls, bachelors, and pregnant cows, to determine maturation capability of the larvae. Hookworms were not recovered from the intestines of 3 pups receiving larvae from belly blubber of bulls, 6 pups receiving larvae from belly blubber of bachelors, and 1 nonexposed pup. Maturation of hookworms did occur in 2 pups exposed to larvae from a mixture of belly blubber, mammary tissue, and milk of pregnant cows. Parasitic 3rd-stage hookworm larvae from belly tissues of pregnant and "non-pregnant" fur seal cows averaged 938.1 and 802.1 micron long, and 34.1 and 31.5 micron wide, respectively; however, larvae from belly tissues of a fur seal bull, bachelors, 2-year-old males, male and female yearlings and pups, and Steller Sea Lion subadults averaged 640.5-732.0 micron long and 20.9-24.9 micron wide.     

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.     

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.     

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.     

Female competition and reproductive success in northern elephant seals

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

Milking strategy in subantarctic fur seals Arctocephalus tropicalis breeding on Amsterdam Island: evidence from changes in milk composition.

Milk composition was investigated throughout the 10-mo pup-rearing period in subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island. The mean milk composition was 42.8% +/- 5.7% lipid, 12.1% +/- 1.5% protein, and 42.6% +/- 7.3% water. Subantarctic fur seals breeding on Amsterdam Island produced one of the richest milks ever reported in otariids (20.4 +/- 2.9 kJ/g), with lipid content contributing 85% of total gross energy. The high lipid levels measured in the milk of subantarctic fur seals breeding on Amsterdam Island is consistent (i) with the relatively long time lactating females spend at sea, due to the relatively poor local trophic conditions near the colony that necessitate that they travel long distances to reach the foraging grounds, and (ii) with the consequently short time mothers spend with their pups ashore. Milk composition changed according to the time mothers were fasting ashore: milk produced during the first 2 d spent ashore, when more than 80% of milk transfer occurred, had higher levels of lipids, proteins, and gross energy than milk produced later during the visit ashore, suggesting that the pups were fed with two types of milk during a suckling period. Throughout the year, mothers in good condition produced milk of higher lipid content than others, suggesting that individual foraging skills contribute to enhance milk quality. Milk lipid and gross energy content varied with pup age, according to quadratic relationships, increasing during the earlier stages of lactation before reaching asymptotic values when pups were 180 d old. The stage of lactation appears to be a better predictor of milk lipid content than the duration of the preceding foraging trip, suggesting that either changes in the nutritional requirements of the pup and/or seasonal changes in trophic conditions act on milk composition. These changes in milk quality may also be related to changes in maternal care; lactating subantarctic fur seals apparently reallocate their body reserves toward gestation rather than lactation at the end of the pup-rearing period.     

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

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

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.     

Influence of maternal mass and condition on energy transfer in Weddell seals

1. Environmental variation influences food abundance and availability, which is reflected in the reproductive success of top predators. We examined maternal expenditure, offspring mass and condition for Weddell seals in 2 years when individuals exhibited marked differences in these traits. 2. For females weighing > or = 355 kg there was a positive relationship between maternal post-partum mass (MPPM) and lactation length, but below this there was no relationship, suggesting that heavier females were able to increase lactation length but lighter females were restricted to a minimum lactation period of 33 days. 3. Overall, females were heavier in 2002, but in 2003 shorter females were lighter than similar-sized females in 2002 suggesting that the effects of environmental variability on foraging success and condition are more pronounced in smaller individuals. 4. There was no relationship between MPPM and pup birth mass, indicating pre-partum investment did not differ between years. However, there was a positive relationship between MPPM and pup mass gain. Mass and energy transfer efficiency were 10.2 and 5.4% higher in 2002 than 2003, which suggests costs associated with a putatively poor-resource year were delayed until lactation. 5. Heavier females lost a higher proportion of mass during lactation in both years, so smaller females may not have been able to provide more to their offspring to wean a pup of similar size to larger females. 6. MPPM had only a small influence on total body lipid; therefore, regardless of mass, females had the same relative body composition. Females with male pups lost a higher percentage of lipid than those with female pups, but by the end of lactation female pups had 4.5% higher lipid content than males. 7. It appears that for Weddell seals the consequences of environmentally induced variation in food availability are manifested in differences in maternal mass and expenditure during lactation. These differences translate to changes in pup mass and condition at weaning with potential consequences for future survival and recruitment.     

Underwater vocalizations and associated behavior in captive ringed seals (<Emphasis Type="Italic">Pusa hispida</Emphasis>)

In pinniped species, especially those that mate in the water, acoustic communication is suggested to play an important role in various aspects of behavior. However, little is known about the behavioral context or function of vocalization, principally because direct observation is difficult in the wild. In the present study, we analyzed the seasonality, sexual differences, and behavioral contexts of the vocalizations of captive ringed seals to explore the function of such communication. The behavior of and underwater sounds made by three ringed seals (an adult male, an adult female, and a subadult female) living in Otaru Aquarium, Japan, were recorded for 19 days between August 2011 and April 2012. Six call types (long snort, knock, yelp, bark, click, and woof) were identified in the recordings. The 12 observed social behaviors could be categorized into three categories (male courtship, aggression, and submission). All call types except clicks were vocalized during social behavior. Vocalizations of all types increased during the breeding season. The long snorts were only produced by the adult male toward an adult female during his courtship behavior. All three individuals emitted knocks, yelps, and bark sounds. Of these three call types, knocks were associated with aggressive behavior or the male’s courtship behavior. In contrast, alternate series of yelps and barks were vocalized by the recipients of aggressive behaviors, suggesting their function as submissive signals. This study could be applied to the monitoring of wild ringed seals with passive acoustic recordings to assess not only their distribution but also their behavior.     

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.     

Vocal Discrimination in Mate Guarding Male Australian Sea Lions: Familiarity Breeds Contempt

The vocal characteristics of a species can be immensely diverse, and can significantly impact animal social interactions. The social structure of a species may vary with geographical variation in call characteristics. The ability of pinnipeds (true seals, fur seals, sea lions and walrus) to distinguish between conspecifics may assist male reproductive strategies, particularly mate acquisition. We assessed the ability of mate guarding Australian sea lion (Neophoca cinerea) males to discriminate local from foreign males’ barks recorded from a geographically distant breeding colony. Bark characteristics were significantly different between colonies, with barks produced by males from the Lewis Island breeding colony higher pitched and longer in both duration and interval duration than barks produced by males on Kangaroo Island. Mate guarding males displayed inter‐colony discrimination of barks, with a significantly stronger response to barks from local males than to those of males from a colony approx. 180 km away. Local males’ barks were apparently considered a greater threat than barks from unfamiliar males. We propose that discrimination of acoustic characteristics may facilitate reproductive isolation in this species that may lead to an ethological–acoustic barrier between breeding colonies, and subsequent genetic isolation.     

The development,inhibition and disinhibition of pup-cannibalism in the Mongolian gerbil

The responses of juvenile and adult gerbils towards newborn test pups were investigated. Young animals of both sexes are less likely to attack the test pup than are older animals. Mature females are more likely to attempt to cannibalize pups than are mature males. Attempted cannibalism by adult males is inhibited when their mate becomes pregnant. If the pregnant female is removed from the cage for one to three days, however, attempted cannibalism is disinhibited. On the other hand, males which have reared a previous litter will not harm test pups when the female is removed. Experience of a pregnant female and of young pups during the pre-weaning stage does not influence the incidence of attempted cannibalism in adulthood. These data suggest that gerbils take full advantage of changing social conditions.     

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.