Ontogeny of body size and shape of Antarctic and subantarctic fur seals

Pre- and post-weaning functional demands on body size and shape of mammals are often in conflict, especially in species where weaning involves a change of habitat. Compared with long lactations, brief lactations are expected to be associated with fast rates of development and attainment of adult traits. We describe allometry and growth for several morphological traits in two closely related fur seal species with large differences in lactation duration at a sympatric site. Longitudinal data were collected from Antarctic (Arctocephalus gazella (Peters, 1875); 120 d lactation) and subantarctic (Arctocephalus tropicalis (Gray, 1872); 300 d lactation) fur seals. Body mass was similar in neonates of both species, but A. gazella neonates were longer, less voluminous, and had larger foreflippers. The species were similar in rate of preweaning growth in body mass, but growth rates of linear variables were faster for A. gazella pups. Consequently, neonatal differences in body shape increased over lactation, and A. gazella pups approached adult body shape faster than did A. tropicalis pups. Our results indicate that preweaning growth is associated with significant changes in body shape, involving the acquisition of a longer, more slender body with larger foreflippers in A. gazella. These differences suggest that A. gazella pups are physically more mature at approximately 100 d of age (close to weaning age) than A. tropicalis pups of the same age

Characterisation of proteins in the milk of fur seals

Milk protein composition was investigated throughout the lactation periods of the Australian fur seal (Arctocephalus pusillus doriferus) and Antarctic fur seal (Arctocephalus gazella). The mean protein content of the milk was found to be 10.9% and 10.6% respectively. The concentration of total protein did not change during lactation, although a decline in casein content of the milk in late lactation was apparent. Milk protein concentration during a foraging/suckling cycle of the Antarctic fur seal analysed at the time of arrival on shore, and 24 h and 72 h after arrival was 12.8%, 11.4% and 12.5% respectively. Re-feeding animals at 72 h resulted in a significant increase in milk protein content to 14.9%. Characterisation of milk protein by SDS-PAGE analysis revealed 5 casein and 10 major whey protein bands. Amino-terminal sequencing indicated that the majority of the whey fraction of the milk is β-lactoglobulin (β-LG). The limited amino acid sequence indicated 3 different β-LGs were secreted in the milk. Subsequently, RT-PCR was used to extend the sequence of one of the β-LGs and translation of the 464 bp fragment indicated that it shared 79% sequence identity with feline β-LG II.

Species-specific characteristics and individual variation of the bark call produced by male Australian fur seals (Arctocephalus pusillus doriferus)

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.

Individual variation in the pup attraction call produced by female Australian fur seals during early lactation

Otariid seals (fur seals and sea lions) are colonial breeders with large numbers of females giving birth on land during a synchronous breeding period. Once pups are born, females alternate between feeding their young ashore and foraging at sea. Upon return, both mother and pup must relocate each other and it is thought to be primarily facilitated by vocal recognition. Vocalizations of thirteen female Australian fur seals (Arctocephalus pusillus doriferus) were recorded during the breeding seasons of December 2000 and 2001, when pups are aged from newborns to one month. The pup attraction call was examined to determine whether females produce individually distinct calls which could be used by pups as a basis for vocal recognition. Potential for individual coding, discriminant function analysis (DFA), and classification and regression tree analysis were used to determine which call features were important in separating individuals. Using the results from all three analyses: F0, MIN F and DUR were considered important in separating individuals. In 76% of cases, the PAC was classified to the correct caller, using DFA, suggesting that there is sufficient stereotypy within individual calls, and sufficient variation between them, to enable vocal recognition by pups of this species.

Variation in maternal provisioning by lactating Antarctic fur seals: response to experimental manipulation in pup demand

An experiment involving the supplementary feeding of pups was conducted on Antarctic fur seals to investigate the factors influencing maternal foraging-attendance cycles and the differential use of nutritional resources for growth, maintenance and storage by pups. For 40% of the lactation period, male pups were given a supplement mimicking the chemical composition of Antarctic fur seal milk at a dose equivalent to 35% of the normal mass-specific milk energy intake for the species. Milk consumption, body composition and growth rates were monitored during and after the supplementary feeding period and maternal foraging-attendance cycles were monitored throughout lactation. During the supplementary feeding period, treatment pups (n=8) grew 32% faster and deposited greater adipose tissue stores than controls (n=8) but consumed the same amount of maternal-delivered milk. When supplementary feeding was stopped (timed to coincide with peak maternal milk yield in this species), treatment pups lost mass whereas control group pups continued to grow. Treatment pups weaned at a younger age (109 days) than control pups (116 days) but at the same mass (13 kg). Maternal attendance durations did not differ between the treatment and control groups throughout lactation. However, mothers of treatment pups had significantly shorter foraging trip durations (3.74 days) than mothers of control pups (4.74 days) during the period of supplementary feeding (there were no significant differences throughout the rest of lactation). These findings are in accordance with predictions of a marginal-value model of fur seal lactation behaviour.

Variation in plasma leptin levels in response to fasting in Antarctic fur seals (Arctocephalus gazella)

Plasma leptin levels were determined in 8 lactating female and 20 pup Antarctic fur seals (Arctocephalus gazella) during fasting periods of normal duration. Plasma leptin levels ranged from 1.35-3.19 ng·ml^-1 in lactating females and 1.79-4.80 ng·ml^-1 in pups and were not positively correlated with body mass or condition. A negative trend, however, was observed between plasma leptin levels and body condition in lactating females upon their arrival at the colony following a foraging trip (beginning of fast). In accordance with findings in other species, plasma leptin levels dropped significantly (P<0.02) in response to the 17-19% drop in body mass experienced by pups during fasting. In contrast, plasma leptin levels in lactating females increased during the first 24 h of fasting before decreasing throughout the remaining 48 h of the fast. This unexpected result could be due to the high level of energy expenditure by seals as they swim back to the colony (i.e. post-exercise response) or may be influenced by the intense suckling activity experienced by females during the onshore fasting periods. The results of this study support recent findings in other carnivore species which suggest the primary physiological role of leptin in these species may not necessarily be as a signal of the magnitude of body energy reserves.

Growth and condition in Australian fur seals Arctocephalus pusillus doriferus (Carnivora : Pinnipedia)

Mass and length growth models were determined for male (n = 69) and female (n = 163) Australian fur seals (Arctocephalus pusillus doriferus) collected at a breeding colony on Seal Rocks (38˚31′S, 145˚06′E), Bass Strait, in south-east Australia, between February and November during 1970–72. Growth was best described by the logistic model in males and the von Bertalanffy model in females. Asymptotic mass and length were 229 kg and 221 cm for males, and 85 kg and 163 cm for females. In all, 95% of asymptotic mass and length were attained by 11 years and 11 years, respectively, in males compared with 9 years and 5 years, respectively, in females. Males grew in length faster than females and experienced a growth spurt in mass coinciding with the onset of puberty (4–5 years). The onset of puberty in females occurs when approximately 86% of asymptotic length is attained. The rate of growth and sexual development in Australian fur seals is similar to (if not faster than) that in the conspecific Cape fur seal (A. p. pusillus), which inhabits the nutrient-rich Benguela current. This suggests that the low marine productivity of Bass Strait may not be cause of the slow rate of recovery of the Australian fur seal population following the severe over-exploitation of the commercial sealing era. Sternal blubber depth was positively correlated in adult animals with a body condition index derived from the residuals of the mass–length relationship (males: r² = 0.38, n = 19, P < 0.001; females: r² = 0.22, n = 92, P < 0.001), confirming the validity of using such indices on otariids. Sternal blubber depth varied significantly with season in adult animals. In males it was lowest in winter and increased during spring prior to the breeding season (r² = 0.39, n = 19, P < 0.03) whereas in females it was greatest during winter (r² = 0.05, n = 122, P< 0.05).

Dive behaviour, foraging locations and maternal-attendance patterns in Australian fur seals (Arctocephalus pusillus doriferus)

The dive behaviour, foraging locations, and colony-attendance patterns of female Australian fur seals (Arctocephalus pusillus doriferus) from Kanowna Island (39°10'S, 146°18'E) in Bass Strait, southeastern Australia, were determined throughout lactation during 1997–1999. Foraging-trip durations increased as lactation progressed, being shortest in summer (3.71 ± 0.24 days; mean ± 1 SE) and longest in winter (6.77 ± 0.57 days, P < 0.05), but maternal-attendance periods did not differ in duration (1.70 ± 0.10 days, P > 0.5). Individual mean attendance periods and trip durations were positively correlated (r² = 0.21, P < 0.005). Diving commenced shortly after seals left the colony (2.6 ± 0.4 h), was continuous for long periods (up to 36 h), occurred mostly during daylight hours, and lacked regular diel variation in depth. The majority of dives (78%) were typically U-shaped and reached depths corresponding to the prevailing depths in Bass Strait (65–85 m), indicating that these animals forage mostly on the benthos of the shallow continental shelf in this region. Such behaviour is unusual for fur seals but is reminiscent of that of some sea lion species. Mean dive durations varied between 2.0 and 3.7 min (maximum 8.9 min) and the theoretical aerobic dive limit (3.91–4.26 min) was exceeded on 17.3% of dives. Dive frequency (8.3 ± 0.6/h) and the proportion of time at sea spent diving (40.7 ± 2.1%) were weakly negatively related to the duration of the foraging trip (r²= 0.07, P < 0.004, and r² = 0.13, P < 0.0001, respectively). Data from at-sea locations showed that lactating females forage almost exclusively within Bass Strait during all seasons.

Interactions between fur seals and a squid jig fishery in southern Australia

Interactions between Australian fur seals (Arctocephalus pusillus doriferus) and the Southern Squid Jig Fishery (SSJF) were investigated in April–May 2002. Details on the number of seals present, distance from the vessel, age and gender, and their behaviour were recorded using scan sampling techniques over 26 nights from eight vessels operating out of Portland and Port Fairy on the southwest coast of Victoria. Seals were observed on all nights but none were recorded in 30% of all (777) scan observations. Of the seals attending vessels at any one time (1.89 ± 0.24), 67% were involved in activities unrelated to jigging operations with the most common behaviour category being resting/grooming. Only 3.6% of observations involved seals targeting squid caught on jig lures whereas a further 29% were of foraging on squid within 40 m of the vessel. Damage to fishing gear attributable to seals was recorded on only three occasions. There was no evidence of negative impacts on seals from vessel operations. The majority of seals foraging on squid around vessels were adult females (71%) with the remainder being almost exclusively juvenile males. The current level of interactions between Australian fur seals and vessels in the SSJF appears minor.

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.

Habitat selection by female Australian fur seals (Arctocephalus pusillus doriferus)

1. Numerous studies have determined the foraging areas of marine apex predators and investigated their relationship to oceanographic features. Most of these, however, have concentrated on surface-feeding seabirds or epipelagic-foraging marine mammals and there is little information on habitat selection in benthic divers.

2. Satellite telemetry was used during the winters of 2001-2003 to determine the foraging areas of 48 female Australian fur seals (Arctocephalus pusillus doriferus) from four breeding sites in northern Bass Strait whose colonies together represent > 80% of the total species population.

3. All individuals foraged over the shallow continental shelf of Bass Strait supporting earlier studies that suggested the species is an exclusively benthic forager. Individual females showed a high degree of foraging site-fidelity and several foraging 'hot spot' areas could be identified.

4. Analysis of habitat use indicated that individuals selected areas with depths of 60-80 m significantly more (λ = 0.216, P<0.001) than any other bathymetric class. There was also evidence for foraging areas being influenced by SST, with individuals selecting regions of 16.0-16.8 C SST (λ = 0.008, P<0.01), but not surface chlorophyll-a concentration (P> 0.05).

5. Temporal analysis of at-sea movements indicated, due to their primarily benthic foraging mode, the areas frequented by female Australian fur seals did not overlap substantially with areas targeted by commercial fisheries. An exception to this was in far eastern Bass Strait where the Otter Trawl component of the Commonwealth Trawl Sector is highly active over the continental shelf and encompasses the areas frequented by females from The Skerries colony.

Temporal structure of diving behaviour in sympatric Antarctic and Subantarctic fur seals

Lactation is considerably briefer (4 vs. 10 mo) and daily pup energy expenditure higher in Antarctic (AFS) than in subantarctic fur seals (SFS), even in sympatric populations of both species, where their foraging locations and diets are similar. Therefore, lactational demands may be higher for AFS females. We investigated whether sympatric lactating AFS and SFS females differ in their physiological or behavioural diving capacities, and in the temporal structure of foraging behaviour. Mean dive depth and duration were greater in SFS, but dives below 140 m were performed only by AFS. An index of activity level during the bottom phase of dives, when fur seals are thought to capture prey, was higher in SFS. Despite these differences, SFS females showed a steady increase in the minimum postdive interval following dives lasting longer than 250 s, compared to the steady increase following dives lasting longer than only 150 s in AFS. These results suggest that physiological constraints on diving behaviour are stronger on AFS females, and that the behavioural aerobic dive limit is greater for SFS. Assuming that dive bouts reflect foraging in prey patches, AFS females exploited more patches per unit time, and remained in them for briefer periods of time, compared to SFS females. Dive bout structure did not differ between overnight and long foraging trips. Our data suggest that AFS females spend greater foraging effort, but may gain access to prey patches of better quality, which may help them cope with higher lactational demands.