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.     

VARIATION IN THE FORAGING LOCATION OF ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) AND THE EFFECTS ON DIVING BEHAVIOR

This study investigated how female Antarctic fur seals adapt their foraging behavior, over time scales of days, to spatial unpredictability in the distribution of their food. Lactating Antarctic fur seals are central-place foragers that feed on highly patchy but spatially and temporally dynamic food. We measured the foraging distribution of 28 fur seals to test whether variation in foraging trip durations was reflected in variation in the location of foraging and the diving behavior of seals at sea. Based on the maximum distance travelled from the breeding beach, three categories of foraging trips were denned: those to the continental shelf area ( n = 12, median = 71 km), to oceanic water ( n = 11, median =164 km), and to farther offshore oceanic waters ( n = 5, median = 260 km). Trip duration and mean surface speed were positively correlated with the maximum distance travelled from the breeding beach. Seals on longer trips spent proportionally less of their time submerged, but there was no significant difference in the total number of dives or the total time spent foraging by seals in relation to trip duration. Evidence from this study and previous work investigating energy gain suggests that an animal on a longer foraging trip could potentially have a higher mean energy return per dive than a similar animal on a shorter foraging trip. Evidence presented suggests that the type of foraging trip (near or far) is not predetermined by the animal but may be a simple response to the stochastic distribution of the resources available.     

BACULAR AND TESTICULAR GROWTH AND ALLOMETRY IN THE CAPE FUR SEAL ARCTOCEPHALUS P. PUSILLUS (OTARIIDAE)

The baculum in Arctocephalus p. pusillus reaches up to 14.1 cm in length, 13.5 g in mass, and 1.3 g/cm in density (= mass/length). A pubertal growth spurt occurs between 2 and 3 yr of age, when bacular length increases by 28%, mass by 124%, and density by 77%; concurrently, body length increases by 14%. A second, weaker spurt occurs at social maturity (9-10 yr of age). Testes grow most rapidly between 1 and 2 yr of age, when testicular length increases by 29%. After 3 yr of age, growth in bacular and testicular length slows, and bacular mass continues to increase approximately linearly. Bacular and testicular lengths average 6.8% and 3.4% (respectively) of body length in adults, compared with 9.9% and 5.7% in the promiscuous harp seal ( Pagophilus groenlandicus ). Bacular length, mass, and density, and testicular length, are positively allometric to body length over growth; bacular length is isometric to testicular length. Among animals of the same age, bacular length and mass are positively allometric to body length in young animals, with negative allometry or isometry thereafter; testicular length is isometric to body length in young animals and negatively allometric thereafter. Patterns of early growth and allometry of the baculum and testes are interpreted as adaptations for mating opportunities, years before territoriality is possible. The baculum and testes of adult Cape fur seals and other otariids are small compared with those of most phocids, because sperm competition among male otariids is weak.     

SUMMER DIVING BEHAVIOR OF MALE WALRUSES IN BRISTOL BAY, ALASKA

Pacific walruses ( Odobenus rosmarus divergens ) make trips from ice or land haul-out sites to forage for benthic prey. We describe dive and trip characteristics from time-depth-recorder data collected over a one-month period during summer from four male Pacific walruses in Bristol Bay, Alaska. Dives were classified into four types. Shallow (4 m), short (2.7 min), square-shaped dives accounted for 11% of trip time, and many were probably associated with traveling. Shallow (2 m) and very short (0.5 min) dives composed only 1% of trip time. Deep (41 m), long (7.2 min), square-shaped dives accounted for 46% of trip time and were undoubtedly associated with benthic foraging. V-shaped dives ranged widely in depth, were of moderate duration (4.7 min), and composed 3% of trip time. These dives may have been associated with navigation or exploration of the seafloor for potential prey habitat. Surface intervals between dives were similar among dive types, and generally lasted 1–2 min. Total foraging time was strongly correlated with trip duration and there was no apparent diel pattern of diving in any dive type among animals. We found no correlation between dive duration and postdive surface interval within dive types, suggesting that diving occurred within aerobic dive limits. Trip duration varied considerably within and among walruses (0.3–9.4 d), and there was evidence that some of the very short trips were unrelated to foraging. Overall, walruses were in the water for 76.6% of the time, of which 60.3% was spent diving.     

INDIVIDUALITY IN THE VOICE OF FUR SEAL FEMALES: AN ANALYSIS STUDY OF THE PUP ATTRACTION CALL IN ARCTOCEPHALUS TROPICALIS

Like most otariids species, the Subantarctic fur seal breeds on land in large, dense colonies. Pups are confronted by the long and repetitive absences of their mother throughout lactation. At each mother's return, pups have to find her among several hundreds of congeners. This recognition process mainly relies on acoustic signals. We performed an acoustic analysis on 125 calls from 20 females recorded during the 1999–2000 breeding season on Amsterdam Island (Indian Ocean). Ten variables were measured in both temporal and frequency domains. To find the acoustic parameters supporting individual signature, we assessed the differences between individuals using Kruskall-Wallis univariate analysis of variance. For each variable, we also calculated the potential of individuality coding (PIC) as the ratio between the between-individual coefficient of variation and the mean value of the within-individual coefficients of variation. We found that the frequency spectrum, the characteristics of the frequency modulation of the initial and middle part of the call and the call duration exhibit an important individual stereotypy (PIC values ranging between 1.5 and 3), whereas features relative to amplitude and the frequency modulation of the final part of the call are weakly individualized (PIC values between 1 and 1.2).     

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.     

REPRODUCTION IN THE FEMALE SUBANTARCTIC FUR SEAL, ARCTOCEPHALUS TROPICALIS

The reproductive tracts of 89 female subantarctic fur seals, taken at Gough Island between November 1977 and October 1978, were examined. Females started ovulating at age 4 yr and all 6-yr-old females were sexually mature. They are mono-ovulatory, alternating between ovaries, and only single embryos were found. Females older than 13 yr ( n = 11) showed poor follicular development and some failed to ovulate. The gestation period (first recorded ovulation to first recorded birth) was 360 d, while delayed implantation (first recorded ovulation to first recorded implantation) lasted for 139 days. Follicle numbers in the functional ovary declined sharply after ovulation while the corpus luteum increased in size until at least 1 mo prior to parturition. The number of follicles in the contralateral ovary increased after midwinter (June/July), and the mean size of the largest follicles peaked prior to ovulation in December. The mean size of the largest follicles increased in both ovaries near implantation time, after reaching a low subsequent to, ovulation. The regressing corpus albicans, conspicuous for approximately 3 mo after parturition, could not be detected macroscopically within one year postpartum. Subantarctic fur seals at Gough Island have a distinct postreproductive class of older females. The pregnancy rate for all females ≥4 yr of age was 79%, and it was 84.5% for the sexually mature group of ≥6 yr of age, while the mean age at puberty was 4.80 yr.     

DIET OF SOUTH AMERICAN FUR SEALS (ARCTOCEPHALUS AUSTRALIS) IN ISLA DE LOBOS, URUGUAY

The diet of Arctocephalus australis was studied through fecal analysis, from September of 1995 to May of 1998, in Isla de Lobos (35°01'50"S-54°53'00"W), Uruguay. A total of 539 scats analyzed indicated that the weakfish ( Cynoscion guatucupa ), the cutlasfish ( Trichiurus lepturus ), the anchoveta ( Engraulis anchoita ), the anchovy ( Anchoa marinii ), and cephalopods were the main prey consumed by these fur seals. Prey richness was higher during January. The diet of A. australis varied between years, apparently related to changes in prey availability. C. guatucupa and E. anchoita decreased while T. lepturus and cephalopods increased from 1996 to 1998. South American fur seals fed on fish ranging in length from 5.4 to 104.8 cm and from 0.7 to 629.9 g in wet mass. However, the mean mass of the main items was never more than 200 g. Observations suggests that A. australis is a trophic generalist with the potential to prey on a wide range of species, although most of the diet is comprised of relatively few species. Fur seals and artisan fisheries both took C. guatucupa , with fur seals taking younger individuals (1–2 yr) than artisan fisheries (> 3 yr).     

INVESTIGATING THE BIASES IN THE USE OF HARD PREY REMAINS TO IDENTIFY DIET COMPOSITION USING ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA) IN CAPTIVE FEEDING TRIALS

The analysis of pinniped scats has been used to quantify their diet, using prey remains to identify species and to estimate the numbers and sizes of prey consumed. There are, however, potential biases involved with scat analysis and, for this method to be used successfully, these biases need to be quantified. Thirty-six Antarctic fur seals ( Arctocephalus gazella ) were fed meals of exclusively either fish, squid, or krill and their scats were collected and analyzed. The major sources of error in the analysis of prey remains from scats were the differential erosion and passage rate of items in relation to their size. However, using simple correction functions, such as those which model otolith erosion, it is possible to reduce these errors. Using plastic beads as dietary markers showed recovery rates were negatively related to their size. Larger squid beaks had lower recovery rates than smaller beaks, but there was no size-related bias in the recovery rates of krill carapaces. Only 33% of the squid beaks and 27% of the otoliths originally fed were recovered from the scats. Recovery rates were greater for squid (77%) and fish (50%) eye lenses and these structures gave a better estimate of numbers eaten. Differences found between experimentally derived and published regression equations (used to calculate prey sizes eaten from prey remains) highlights the need for regression equations based on local prey characteristics, if these are to be used with any success to describe the prey eaten.     

DEVELOPMENT OF HEMOGLOBIN, HEMATOCRIT, AND ERYTHROCYTE VALUES IN GALÁPAGOS FUR SEALS

We studied the ontogeny of hemoglobin concentration, hematocrit and erythrocyte counts in the Galapagos fur seal ( Arctocephalus galapagoensis , Heller 1904). Two hundred and fifty-three animals were sampled between the ages of 22 d and > 8 yr, of which 46 were adult females. Body mass increased steadily with age from 6.1 ± 1.2 kg in 1-mo-old pups ( n = 27) to 28.5 ± 3.3 kg in adult females. Even adult females increased in mass with age. Hemoglobin (Hb), hematocrit (Hct), and red blood cell (RBC) values all increased in a logarithmic fashion with age up to 2 yr. Blood values for pups were Hct: 35.5 ± 4.1%; Hb: 12.9 ± 1.3 g/dl; RBC: 4.1 ± 0.3·10⁶/μl. Half-year-old fur seals (Hct: 42.1 ± 3.2%; Hb: 15.7 ± 1.3 g/dl; RBC: 4.9 ± 0.5·10⁶/μl; n = 50) were the oldest age group to show significantly lower blood values than adult females ( P < 0.001 for all three parameters). Yearling blood values (Hct: 47.2 ± 3.6%; Hb: 17.3 ± 1.6 g/dl; RBC: 5.6 ± 0.4·10⁶/μl; n = 56) did not differ significantly from those of adult females ( P < 0.32; P < 0.26; P < 0.23, respectively). Blood values of adult females were lower than those of 2-yr-olds (Hct: 49.6 ± 2.4%; Hb: 18.5 ± 1.2 g/dl; RBC: 5.7 ± 0.3·10⁶/μl; n = 31). These differences were significant only for RBCs ( P < 0.003). Up to the age of 1 yr, age was the best predictor for blood values, thereafter mass tended to be a better predictor. Female juveniles between the ages of 150 and 600 d had higher blood values than same-age males. The relationship of blood value development to diving activity is briefly described and the results are compared to values of other marine mammals. Ontogeny is discussed in relation to the development of these blood values in terrestrial mammals.     

DIVING BEHAVIOR OF THE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDII) IN MONTEREY BAY, CALIFORNIA

Physical environment and physiological characteristics of marine mammals potentially affect the duration and depth of diving. Härkönen (1987b) proposed a hypothesis that the harbor seal would gain maximum energy by foraging at intermediate depths. To investigate this hypothesis, we studied diving behavior of the Pacific harbor seal (Phoca vitulina ricbardii) during 1995 through 1997 in Monterey Bay, California. Dive depths (n = 13,063 dives) were recorded via time‐depth recorders. Approximately 80% of recorded dives were classified as square dives (type I), which typically were associated with foraging in pinnipeds. Approximately 11% of dives were V dives (type II; 1,402 dives), and the remainder (1,225 dives) were skewed dives (type III and IV). The deepest recorded dive was 481 m, while the greatest duration was 35.25 min. Body mass explained the variability of durations of long dives for females (95th percentile; D₉₅♂=‐5.47 + 0.18 × (mass♀), r²= 0.91, 95% CI for slope = [0.08, 0.28], n= 5) and for males (D₉₅♂=‐5.86 + 0.18 × (mass♀), r²= 0.83, 95% CI for slope = [0.12, 0.24], n= 11). The large proportion of variability in deep dives, however, was explained by body mass only for males (95th percentile; Z₉₅♂=‐363.9 + 6.05 × (mass♀), r²= 0.83, 95% CI for slope = [3.93, 8.17], n= 11) and not for females (Z₉₅,♂=?148.1 +3.11 × (mass♀), r²= 0.58, 95% CI for slope = [‐1.7, 7.9], n= 5, 95% CI for slope= [?1.7, 7.9]). Median depths of presumed foraging dives of harbor seals in the Monterey Bay area were between 5 and 100 m, which were within the range of the previously reported depths for other areas (< 100 m). Our findings generally supported Härkönen's hypothesis that harbor seals forage in the intermediate depth in their environment.     

DIVING BEHAVIOR OF SUBADULT AND ADULT HARBOR SEALS IN PRINCE WILLIAM SOUND, ALASKA

Satellite-linked depth recorders (SDRs) were attached to 47 harbor seals in Prince William Sound, Alaska, during 1992–1996. Parameters describing diving effort, diving focus, and focal depth (depth bin to which diving was focused) were calculated from binned data on maximum dive depth and time spent at depth, and analyzed using repeated-measures mixed models. This analysis method accounted for individual variability, temporal autocorrelation, and the binned nature of SDR data, which are often ignored using standard statistical techniques. Results indicated that diving effort remained steady from September to April, when seals spent 68%-75% of their overall time in the water. Time spent in the water declined to 60% in May and to about 40% in July. Seals spent the most time in the water at night and the least in the morning. The diving of all seals in all months was highly focused. Overall, diving was focused to one depth bin approximately 75% of the time. Diving was more focused for females than for males and subadults. Focal dive depth was deepest in winter and shallowest during May-July. Focal depth and diving focus varied by region. Collinearity between month and region in the focal depth model suggests that seals move in winter to regions where prey are found deeper in the water column. Variations in diving behavior presumably result from combinations of regional bathymetry, seasonal cycles in type or depth distribution of prey, and seal life-cycle events such as reproduction and molting.     

A KINEMATIC STUDY OF SUCTION FEEDING AND ASSOCIATED BEHAVIOR IN THE LONG-FINNED PILOT WHALE, GLOBICEPHALA MELAS (TRAILL)

Analysis of videotaped feeding sequences provides novel documentation of suction feeding in captive juvenile long-finned pilot whales ( Globicephala melas ). Swimming and stationary whales were videotaped while feeding at the surface, mid-water, and bottom. The ingestion sequence includes a preparatory phase with partial gape followed by jaw opening and rapid hyoid depression to suck in prey at a mean distance of 14 cm (duration 90 msec), although prey were taken from much greater distances. Depression and retraction of the large, piston-like tongue generate negative intraoral pressures for prey capture and ingestion. Food was normally ingested without grasping by teeth yet was manipulated with lingual, hyoid, and mandibular movement for realignment; suction was then used to transport prey into the oropharynx. Whales frequently rolled or inverted before taking prey, presumably to avoid grasping and repositioning. Prey were sucked off the bottom or sides of the pool without direct contact; lateral suction was used to ingest items from the sides of the mouth.