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.     

STUDY OF VITAMIN A SUPPLEMENTATION IN CAPTIVE NORTHERN FUR SEALS (CALLORHINUS URSINUS) AND ITS EFFECT ON SERUM VITAMIN E

Two experiments were conducted to determine the effect of vitamin A supplementation on serum vitamin E in adult female northern fur seals ( Callorhinus ursinus ). In the first experiment five animals received, in addition to their routine dietary multivitamin supplement, a high-level vitamin A supplement (53 μmol retinyl palmitate/d) for 30 d. Five seals consuming their routine dietary supplement served as controls. Serum vitamin E decreased significantly in animals receiving high-level vitamin A supplements. At the end of 30 d serum vitamin E averaged 18.6 μg/mL in the control animals and 13.4 μg/mL in the animals receiving the high-Ieve1 vitamin A supplement. In experiment 2 ten animals received the high level vitamin A supplement for 60 d. After 30 d, serum vitamin E levels were reduced, but by 60 d had returned to baseline levels. However, the ratio of serum vitamin E to phospholipid, another index of vitamin E status, remained decreased. Although the exact mechanism of interaction is unknown, this study shows that when providing vitamin supplements for captive pinnipeds, vitamin interactions must be considered. The vitamin A supplementation currently used by some institutions seems unnecessary and may have detrimental effects on vitamin E status.     

DENSITY DEPENDENCE IN NORTHERN FUR SEALS (CALLORHINUS URSINUS)

This paper summarizes, updates, and interprets information on density-dependent dynamics of populations of the northern fur seal ( Callorhinus ursinus ). Density-dependent changes observed in these populations have involved various aspects of growth (body length, body weight, tooth weight, and size of other skeletal parts, especially the skull), survival, age at maturation, incidence of disease, and time spent foraging. For the population of northern fur seals on St. Paul Island of the Pribilof Islands, which was observed during a major increase and during two significant declines, density-dependent changes that occurred during the growth of the population were reversed during the declines. The Robben Island population in the western Pacific declined after 1965, and the decline was accompanied by changes similar to those observed during the declines on the Pribilof Islands. Although data are not available for all age and sex classes, it appears that most or all components of the populations exhibit similar changes. The overall implication of these changes is that current populations on Robben Island and the Pribilof Islands are reduced to levels below what could be supported by the resources available in their environments. Density dependence for this species is consistent with that of other large mammals, specifically in that vital rates for fur seals ate related to density in a nonlinear fashion.     

AUDIOMETRIC ASSESSMENT OF NORTHERN FUR SEALS, CALLORHINUS URSINUS

Aerial and underwater audiograms for two young female northern fur seals ( Callorhinus ursinus ) and one young female California sea lion (Zalophus californianus) were obtained with the same procedure and apparatus. Callorhinus hears over a larger frequency range and is more sensitive to airborne sounds than Zalophus or any other pinniped thus far tested in the frequency range of 500 Hz to 32 kHz. Sensitivity of Callorhinus to waterborne pure tones, ranging from 2 to 28 kHz, is equal or superior to all other pinnipeds tested in this same frequency range. Like Zalophus , the upper frequency limit for underwater hearing (as defined by Masterton et al. 1969) in Callorhinus is about one-half octave lower than the three phocid species thus far tested. Callorhinus' upper frequency limit in air is about 36 kHz and under water it is about 40 kHz. Comparison of air and water audiograms shows Callorhinus is no exception to previous behavioral findings demonstrating that the „pinniped ear” is more suitable for hearing in water than in air. Similar to Zalophus and Phoca vitulina, Callorhinus shows an anomalous hearing loss at 4 kHz in air. The basis for this insensitivity to airborne sounds at 4kHz and not at lower or higher frequencies is presumably caused by specialized middle ear mechanisms matching impedance for waterborne sounds. Critical ratio curves for Callorhinus are similarly shaped to ones obtained for humans but are shifted upwards in frequency. Compared to all other marine mammals thus far evaluated, the critical ratios for Callorhinus are the smallest yet reported.     

MAXIMUM NET PRODUCTIVITY LEVEL ESTIMATION FOR THE NORTHERN FUR SEAL (CALLORHINUS URSINUS) POPULATION OF ST. PAUL ISLAND, ALASKA

The goal of this study was to assess the maximum net productivity level for the northern fur seal ( Callorhinus ursinus ) population of St. Paul Island, Alaska. Definitive determination of this level is not possible due to uncertainty in life table parameters and density-dependent changes in those parameters. To account for such uncertainty, repetitive numerical simulations were used to generate frequency distributions of estimates for the maximum net productivity level and related population parameters. This approach systematically varied simulation input parameters, ran a separate simulation with each input parameter combination, and validated the simulations on the basis of comparison with historical observations. Results from validated simulations were compiled in frequency distributions to provide a measure of confidence for MNPL estimates. The distributions confirm that this population is probably well below its maximum net productivity level. Because they reflect the uncertainty in our understanding of northern fur seal population dynamics, these distributions are a more realistic basis for management than single point estimates.     

TOOTH WEIGHTS OF JUVENILE MALE NORTHERN FUR SEALS, CALLORHINUS URSINUS

Tooth weight and body length of juvenile male northern fur seals, Callorhinus ursinus , collected on St. Paul Island, Alaska, from 1948 to 1984 were compared. The relationship between these variables was investigated for individual and combined age classes. A density-dependent increase in mean tooth weight occurred during a decline in fur seal abundance. Consistent, though poorly understood, differences in tooth weight were found among males associated with different rookeries (breeding grounds). Storing the teeth for long periods may affect the weights, but such effects, if any, are too small to account for the changes observed.     

REPRODUCTIVE SYNCHRONY AND THE ESTIMATION OF MEAN DATE OF BIRTH FROM DAILY COUNTS OF NORTHERN FUR SEAL PUPS

Two methods for estimating the mean date of birth from daily counts of northern fur seal pups ( Callorhinus ursinus ) are presented and applied to data collected on the Pribilof Islands in 1951, 1962, 1963 and 1983. The mean date of birth over the four years was July 9. Reproduction is highly synchronized and consistent from one year to the next. Pupping occurs over a five week period with over 50% of the pups being born during the first two weeks of July.     

COMPARATIVE POPULATION DYNAMICS OF FUR SEALS

The population sizes, trends, exploitation, and life history parameters for the ten fur seal species and subspecies are summarized. The largest population is that of Arctocephalus pusillus pusillus with approximately two million seals, and the smallest is A. townsendi with approximately 7,000 individuals. Most populations are legally protected, although controlled harvesting may occur. None of the fur seal populations is currently known to be decreasing. Data are presented for parameters related to the survival of pups, juveniles, adults, and territorial males, and to reproduction, including the age of attainment of territorial status, aggregation sizes, age of first parturition, pregnancy rates, sex ratios of young animals, and information on the birth seasons of the different species. Since pinnipeds are often of concern in fisheries management, their daily consumption rates are of importance, and consequently data on body masses are summarized and the paucity of data on consumption rates as a function of body mass noted. A simplified age-structured model is developed, and the results of this model are compared with results from more detailed models based on two published life tables for Callorhinus ursinus. This comparison shows that the use of the simplified age-structured model is justified to explore changes in population growth rate. However, the simplified model does show exaggerated age structure effects compared to the more detailed models. This model is used to compare the population dynamics of those species for which sufficient data are available. Areas in which limited, or no, data are available for the different fur seal species are highlighted.     

THE INFLUENCE OF CLIMATIC SEASONALITY ON THE LIFE CYCLE OF THE PRIBILOF NORTHERN FUR SEAL

Weather conditions recorded from 1956 to 1986 on St. Paul Island, Alaska, were probed to establish their influence upon the northern fur seal's life cycle ( Callorhinus ursinus ). Air temperatures, wind speeds, and relative humidity levels were seasonally decomposed and compared with the timing of pupping and migration. Most pups are born in early July when air temperatures and relative humidity approach their highest annual levels and wind speeds are at their lowest. Weather conditions favor growth and survival of pups from July to September but are unfavorable in June. A rapid deterioration in weather through October and November corresponds with the fall migration of pups and lactating females. The data suggest the pivotal event in the fur seal's life cycle is the timing of birth and survival of nursing pups. As such, the ultimate determinant of the precisely timed fur seal life cycle appears to be climatic seasonality during the breeding season.     

GENERIC NAMES OF NORTHERN AND SOUTHERN FUR SEALS (MAMMALIA: OTARIIDAE)

We have resolved a nomenclatural problem discovered during research on the northern fur seal that concerns the correct generic name for this taxon and for fur seals of the Southern Hemisphere. The unfortunate practice by some 19th-century authors to use names in their Latinized form but to date them from their first appearance as French common names led to the use of Arctocepbalus for southern fur seals when the name correctly applies to the northern fur seal, known today as Callorbinus ursinus . However, Arctocepbalus and Callorbinus are antedated by Otoes G. Fischer, 1817, which is the earliest available generic name for the fur seal of the northern Pacific. The earliest available generic name for southern fur seals is Halarctus Gill, 1866. To avoid the confusion that would result from replacing the currently used generic names with those required by strict adherence to the Principle of Priority, we have petitioned the International Commission on Zoological Nomenclature to preserve Arctocepbalus and Callorbinus for the southern and northern fur seals, respectively.     

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.     

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.     

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.     

INVESTIGATING TROPHIC RELATIONSHIPS OF PINNIPEDS IN ALASKA AND WASHINGTON USING STABLE ISOTOPE RATIOS OF NITROGEN AND CARBON

We measured stable-nitrogen (δ¹⁵N) and stable-carbon (δ¹³C) isotope ratios in muscle and hair from 7 northern fur seals (Callorhinus ursinus) from the Pribilof Islands, Alaska, and 27 Steller sea lions (Eumetopias jubatus), and 14 harbor seals (Phoca vitulina) from the Gulf of Alaska and coast of Washington State, in order to contrast dietary information derived from isotopic vs. available conventional dietary studies. Stable-nitrogen-isotope analysis of muscle revealed that harbor seals were enriched over sea lions (mean δ¹⁵N = 18.6‰vs. 17.5‰) which were in turn enriched over northern fur seals (mean δ¹⁵N = 16.6‰). Trophic segregation among these species likely results primarily from differential reliance on herring (Clupea harengus), Atka mackerel (Pleurogrammus monopterygius), and large vs. small walleye pollock (Theregra chalcogramma). According to their δ¹⁵N values, adult male Steller sea lions showed a higher trophic position than adult females (mean δ¹⁵N: 18.0‰vs. 17.2‰), whereas adult female northern fur seals were trophically higher than juvenile male fur seals (mean δ¹⁵N: 16.5‰vs. 15.0‰). Each of these observed differences likely resulted from differential reliance on squid or differences in the size range of pollock consumed. Three northern fur seal pups showed higher δ¹⁵N enrichment over adults (mean 17.7‰vs. 15.8‰) due to their reliance on their mother's milk. Stable-carbon isotope measurements of hair revealed a cline toward more negative values with latitude. Segregation in hair δ¹³C between Steller sea lions and harbor seals off the coast of Washington (mean δ¹³C: ?13.6‰vs.?15.0‰) reflected the greater association of harbor seals with freshwater input from the Columbia River. Our study demonstrates the utility of the stable isotope approach to augment conventional dietary analyses of pinnipeds and other marine mammals.     

INDICES OF BODY CONDITION AND BODY COMPOSITION IN FEMALE ANTARCTIC FUR SEALS (ARCTOCEPHALUS GAZELLA)

An attempt was made to develop simple, inexpensive, rapid means of determining body composition in Antarctic fur seals ( Arctocephalus gazella ). Measurements of total body water ( TBW ) and total body lipid ( TBL ), obtained by hydrogen isotope dilution, were compared to the results of bioelectrical impedance analysis ( BIA ) and morphometric indices of body condition in 52 adult females. TBW was weakly correlated with BIA measurements of resistance ( v = -0.30, P < 0.03). Conductor volume (length2/resistance) was more highly correlated with TBW ( r = 0.75, P < 0.0001) and the inclusion of mass into the predictive equation improved the correlation further ( r = 0.95, P < 0.0001). A body condition index (mass/length) previously used in pinniped studies was positively correlated to TBL ( r = 0.77, P < 0.0001) validating its use as a relative index of condition. However, body mass alone was highly correlated to TBW ( r = 0.94, P < 0.0001) and appears to provide a simple, rapid means of estimating body composition in adult females. This technique may also be applicable to juvenile male Antarctic fur seals.     

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).     

DIET OF RINGED SEALS (PHOCA HISPIDA) ON THE EAST AND WEST SIDES OF THE NORTH WATER POLYNYA, NORTHERN BAFFIN BAY

In conjunction with the International North Water Polynya Study (NOW) in northern Baffin Bay, we examined the diets of ringed seals ( Phoca hispida )¹ on the west (Grise Fiord, Nunavut) and east (Qaanaaq, Greenland) sides of the polynya, using conventional stomach content analysis, as well as inferences from stable isotope ratios in seal muscle. Between May and July 1998, stomach and muscle tissue samples were collected from 99 ringed seals taken near Grise Fiord and 100 taken near Qaanaaq. The amphipod Themisto libellula was the dominant prey type in the diet of immature ringed seals from Grise Fiord, whereas arctic cod ( Boreogadus saida ) and polar cod ( Arctogadus glacialis ) predominated in the diet of adults. Both immature and adult seals collected near Qaanaaq fed predominantly on arctic cod. Overall, seals collected near Grise Fiord had significantly higher δ¹³C values than those collected near Qaanaaq ( P < 0.001), but there was no statistical separation in δ¹⁵N values between the two samples ( P = 0.06). Differences in diets of ringed seals from the east and west sides of the North Water Polynya may be due to differences in prey distribution and/or differences in biological productivity and fish biomass within the polynya.