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.     

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.     

ANALYSIS OF SWIM VELOCITIES DURING DEEP AND SHALLOW DIVES OF TWO NORTHERN FUR SEALS, CALLORHINUS URSINUS

Swim velocities at 15-sec intervals and maximum depth per dive were recorded by microprocessor units on two "mixed diver" adult female northern fur seals during summer foraging trips. These records allowed comparison of swim velocities of deep (>75 m) and shallow (<75 m) dives.
Deep dives averaged 120 m depth and 3 min duration; shallow dives averaged 30 m and 1.2 min. Mean swim velocities on deep dives were 1.8 and 1.5 m/sec for the two animals; mean swim velocities on shallow dives were 1.5 and 1.2 m/sec. The number of minutes per hour spent diving during the deep and shallow dive patterns were 11 and 27 min, respectively.
Swim velocity, and hence, relative metabolic rate, did not account for the differences in dive durations between deep and shallow dives. The long surface durations associated with deep dives, and estimates of metabolic rates for the observed swim velocities, suggest that deep dives involve significant anaerobic metabolism.     

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.     

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.     

MOVEMENTS OF SOUTHERN ELEPHANT SEALS AND SUBANTARCTIC FUR SEALS IN RELATION TO MARION ISLAND

Based primarily on an intensive marking/resighting program conducted at Marion Island (46°54'S, 37°45'E) in the Southern Ocean, the inter-island movements of southern elephant seals, Mirounga leonina , and fur seals Arctocephalus spp., were investigated to elucidate their little known pelagic phase. Southern elephant seals, in particular immature animals, readily move between the proximate Marion and Prince Edward islands. Some range as far afield as Iles Crozet, approximately 1,000 km distant where they haul out for the summer molt or during an autumn resting phase. The exchange of individuals between Marion Island and Iles Crozet during the return of immatures for the molt after a winter at sea, suggests overlapping of the foraging ranges of the two populations. Despite their wanderings, the majority of M. leonina from Marion Island probably feed in the proximity of the island, and relocate onto the island for breeding, molting and resting. Of the fur seals, only a few A. tropicalis were seen away from their natal island, in some cases covering distances in excess of 2,000 km. displaying a remarkable dispersal capacity.     

ENTANGLEMENT IN MAN-MADE DEBRIS OF ANTARCTIC FUR SEALS AT BIRD ISLAND, SOUTH GEORGIA

Neck collars of man-made marine debris were seen on 208 Antarctic fur seals (and removed from 170) during the 142 days of the 1988–1989 pup-rearing season at Bird Island, South Georgia. This represents at least 0.1% of the total Bird Island population and a minimum of 0.4% of animals in the best covered areas; a maximum value might approach 1%. Polypropylene straps (packaging bands) formed 59% of collars, nylon string (16%), fishing net (13%) and six other materials comprised the rest. Males accounted for 71% of entanglements, 88% of which were of young (1–4 yr old) animals; females accounted for 64% of animals older than this. Obvious physical injury was being caused to 30% of animals and only on 19% of animals was the collar loose enough potentially to come off. The magnitude of the problem at South Georgia is similar to that with northern fur seals at the Pribilof Islands, where a significant population decline has occurred concurrently. Antarctic fur seals are still increasing in numbers but stricter controls on the jettisoning of debris into the Southern Ocean ate needed if the entanglement problem is not to increase beyond the level of a potential threat.     

EVALUATION OF AN EFFECTIVE METHOD TO ESTIMATE AGE OF CAPE FUR SEALS USING GROUND TOOTH SECTIONS

Teeth of known-age Cape fur seals Arctocephalus pusillus pusillus were used to validate age estimated from ground sections. In the canines, dentine growth layer groups (GLGs) reflected age accurately but no reliable readings could be obtained from GLGs in the cementum. Upper canines were the most suitable for age estimation. By contrast, in the postcanines where the cementum is thicker, only GLGs in the cementum could be used for age determination, but not with the same accuracy as for dentine in the canines. Therefore, it is recommended that GLGs in the dentine be used to determine age in the Cape fur seal. However, pulp cavities in canines closed at about 13 yr and consequently GLGs in the cementum of the postcanines should be used where the pulp cavities of canines are closed. Accurate estimation of age is not possible from the dentine of older animals.     

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.     

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.     

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.     

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.     

SUCCESSFUL USE OF A TRANSLOCATION PROGRAM TO INVESTIGATE DIVING BEHAVIOR IN A MALE AUSTRALIAN FUR SEAL, ARCTOCEPHALUS PUSILLUS DORIFERUS

This study reports some of the first foraging behavior data collected for male fur seals. A nonbreeding male Australian fur seal, Arctocephalus pusillus doriferus , captured at a commercial salmon farm in southern Tasmania, Australia, was relocated 450 km from the site of capture. The animal was equipped with a geolocating time-depth recorder that recorded diving behavior and approximate location for the 14.4 d that it took the seal to travel down the east coast of Tasmania and be recaptured at the salmon farm. During its time at sea, the seal spent most of its time over the relatively shallow shelf waters. It spent 30% of its time ashore on a number of different haul-out sites. The deepest dive was 102 m and the maximum duration was 6.8 min. "Foraging" type dives made up 31.2% of the time at sea and had a median duration of 2.5 min and a median depth of 14 m. The seal performed these dives more commonly during the latter part of its time at sea, while it was on the east coast. Unlike other fur seal species studied to date, there was no evidence of a diurnal foraging pattern; it made dives at all times of the day and night.     

Early development, adult mass, and reproductive success in bighorn sheep

Despite considerable empirical and theoretical work on the individualand population consequences of early development, little isknown about the correlations between early mass and adult sizeor lifetime reproductive success of free-ranging mammals. Usinga 26-year study of bighorn sheep (Ovis canadensis), we examinedhow mass as a lamb and mass gain as a yearling affected adultmass for both sexes, horn length of males and lifetime reproductivesuccess of females at different population densities. Mass asa 3-week-old lamb was either weakly or not correlated withadult mass, horn length of adult males, or the number of lambsweaned over a ewe's lifetime. Weaning mass was correlated withmost of these variables when the number of ewes in the populationwas taken into account. When weaning mass was controlled throughpartial correlation, mass as a yearling was correlated with adult mass of ewes but not with ewe reproductive success orwith adult mass or horn length of rams. Lamb mass and numberof ewes explained more of the variance in adult characteristicsfor males than for females. Our results suggest that mass gainduring lactation, possibly but not necessarily related to theamount of maternal care received, affects adult mass and reproductive success. Females appear better able than males to compensatefor poor early development, likely by postponing their firstreproduction. Mass gain over several years and the number ofewes in the population strongly affect adult mass of both sexesand therefore can have profound effects on reproductive successof this long-lived species with a multi-year growth period.