GENETIC SUBSTRUCTURE OF THE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDSI) OFF WASHINGTON, OREGON, AND CALIFORNIA

Genetic substructure among groups of Pacific harbor seals, Phoca vitulina richardsi , along the western coast of the United States was investigated using mitochondrial DNA sequences. Blood and tissue samples were removed from 86 seals inhabiting Puget Sound and the Pacific coasts of Washington, Oregon, and California. A 320 base-pair segment of the control region was amplified using the polymerase chain reaction and directly sequenced. These data indicated a high level of diversity. Thirty variable sites were found that define 47 mitochondrial haplotypes. Among groups of P. v. richardsi sampled, 5 haplotypes were shared, but most (42) were unique to a locality. Haplotypic frequency and an Analysis of Molecular Variance (A mova ) revealed significant differences ( P = 0.001) among regions. Phylogenetic analysis indicated Puget Sound seals possess unique divergent lineages not found in seals from the coasts of Washington, Oregon, and California. These lineages may represent haplotypes from north of Washington, which is consistent with late reproductive timing of harbor seals from Puget Sound.     

PUPPING PHENOLOGY AND DEMOGRAPHY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI)ON TUGIDAK ISLAND, ALASKA

The number of seals on shore at Tugidak Island (Gulf of Alaska) declined 72%–85% between 1976 and 1988 and increased during the 1990s. We compared pupping phenology and the ratio of pupping-period counts to molting-period counts between declining (1976–1979) and increasing (1994–1998) years, and examined the sex/age structure of seals ashore during the 1990s. In the 1970s the onset and peak of pupping occurred 6–18 d later than in the 1990s. Rate of pup abandonment was higher in 1978 than in the 1990s. Between 1994 and 1995, the maximum and mean number of seals ashore increased >50%, largely due to an increase in non-pups. From 1995 to 1998, the sex/age structure of seals ashore was similar among years. We observed three to four times as many seals during the molting period than during the pupping period in the 1970s, whereas similar numbers were ashore during these periods in the 1990s, perhaps reflecting changes in demography and/or haul-out behavior. Changes in pupping phenology and demography may reflect environmental changes, such as food availability, and when monitored in conjunction with population counts, may help us better interpret changes in the number of seals ashore.     

A REASSESSMENT OF THE IMPACT OF THE EXXON VALDEZ OIL SPILL ON HARBOR SEALS (PHOCA VITULINA RICHARDSI) IN PRINCE WILLIAM SOUND, ALASKA

Analyses of population trends and movements of harbor seals in Prince William Sound (PWS) casts doubt on published findings that 302 seals were killed by the Exxon Valdez oil spill in 1989. Assumptions that seals have 100% fidelity to a haul-out, that they were not displaced by the spill and associated disturbances, and that population trends throughout PWS varied similarly, except for oil spill effects, are not supported. Survey efforts to account for missing seals in 1989 were incomplete, too late in the year, and geographically limited. Basic assumptions required for statistical comparisons of oiled and unoiled haul-outs were violated. Fourteen dead seals, mostly pups, were recovered in PWS. Cause of death in most instances could not be determined, nor could the proportion that would have died naturally. Evidence does not support high unsubstantiated mortality, but is more consistent with seals avoiding or moving away from some oiled haul-outs. Interpretation of survey results requires consideration of temporal and regional variation. "Route A" surveys of central and eastern PWS do not represent population trends in western PWS or at glacial haul-outs. To adequately monitor population trends of PWS as a whole, broader sampling must be conducted on a routine basis.     

EVALUATION OF THE ALASKA HARBOR SEAL (PHOCA VITULINA) POPULATION SURVEY: A SIMULATION STUDY

We used simulation to investigate robust designs and analyses for detecting trends from population surveys of Alaska harbor seals. We employed an operating model approach, creating simulated harbor seal population dynamics and haul-out behavior that incorporated factors thought to potentially affect the performance of aerial surveys. The factors included the number of years, the number of haul-out sites in an area, the number and timing of surveys within a year, known and unknown covariates affecting haul-out behavior, substrate effects, movement among substrates, and variability in survey and population parameters. We found estimates of population trend were robust to the majority of potentially confounding factors, and that adjusting counts for the effects of covariates was both possible and beneficial. The use of mean or maximum counts by site without covariate correction can lead to substantial bias and low power in trend determination. For covariate-corrected trend estimates, there was minimal bias and loss of accuracy was negligible when surveys were conducted 20 d before or after peak haul-out attendance, survey date became progressively earlier across years, and peak attendance fluctuated across years. Trend estimates were severely biased when the effect of an unknown covariate resulted in a long-term trend in the fraction of the population hauled out. A key factor governing the robustness and power of harbor seal population surveys is intersite variability in trend. This factor is well understood for sites within the Prince William Sound and Kodiak trend routes for which at least 10 consecutive annual surveys have been conducted, but additional annual counts are needed for other areas. The operating model approach proved to be an effective means of evaluating these surveys and should be used to evaluate other marine mammal survey designs.     

PUPPING IN THE MOST NORTHERLY HARBOR SEAL (PHOCA VITULINA)

The Norwegian high arctic archipelago Svalbard is the home of the world's northernmost population of harbor seals. Due to their limited distribution in isolated areas of the archipelago, little is known about their biology. Until now no information has been published on their pupping or whether it differs from that of harbor seals in other regions. This paper shows that the peak pupping period for harbor seals in Svalbard coincides with that in Europe, i.e., the second half of June. The pups are born with an adult-like pelt and not with a white coat.     

IMMUNOLOGICAL AND HEMATOLOGICAL PARAMETERS IN CAPTIVE HARBOR SEALS (PHOCA VITULINA)

Peripheral blood was collected from 13 captive seals (12 harbor seals and 1 gray seal) of various ages to study different immunological and hematological parameters. In vitro mitogenic reactivity of blood lymphocytes was measured by means of a microculture lymphocyte transformation test (LTT). After stimulation with different doses of the mitogens concanavalin A (Con A) and pokeweed mitogen (PWM), all examined seals showed significant proliferative responses to each mitogen. Furthermore, mitogenic reactivity significantly decreased with animal age suggesting this parameter of seal lymphocyte function is age-related. The present experiments support that the LTT is a suitable tool to monitor the functional capacity of seal lymphocytes. By means of the erythrocyte-rosette (E:-rosette) test it was demonstrated that a subpopulation of mononuclear seal bloosd leucocytes formed rosettes with sheep red blood cells (Srbc). This observation indicates that the phenomenon probably represents a marker for T lymphocytes in the seal as in several other mammalian species. Furthermore, the percentage of Srbc rosette-forming cells decreased with the age of the animals. Total blood leucocyte counts and differential leucocyte counts were determined by light microscopy. The number of leucocytes varied considerably among individual animals. Parallel to a significant decline of the percentage of lymphocytes with age, a corresponding increase in the percentage of neutrophils was demonstrated. In all of the seals, the percentages of monocytes and eosinophils were found to be low with only minor individual variation. The LTT and the E-rosette test are suggested as potential tools to elucidate immunological disorders in the seal.     

ADRENAL FUNCTION IN WILD AND REHABILITATED PACIFIC HARBOR SEALS (PHOCA VITULINA RICHARDII) AND IN SEALS WITH PHOCINE HERPESVIRUS-ASSOCIATED ADRENAL NECROSIS

Adrenal function in harbor seals (Phoca vitulina richardii) was evaluated using adrenocorticotrophic hormone (ACTH) stimulation tests and fecal cortisol levels. The effect of ACTH administration on plasma cortisol and aldosterone levels in five free-living and 14 rehabilitated harbor seal pups was determined using enzyme immunoassay and radioimmunoassay, respectively. In free-living seals, injection of ACTH caused a significant increase in mean plasma cortisol but not of mean aldosterone levels 60 min postinjection. In these seals, mean initial plasma aldosterone was significantly higher than initial levels in rehabilitated seals, while initial cortisol levels were similar. Of the rehabilitated seals, eight died with adrenal cortical necrosis associated with herpesvirus inclusions, while six lived to be released. In the seals that were released, both mean initial cortisol levels and response to ACTH decreased through rehabilitation. In the seals that died, mean initial cortisol and response to ACTH increased through rehabilitation. The differences between initial cortisol levels in seals that lived and those that died were significant at weeks two and four of rehabilitation but not at the week of admission. There was considerable individual variation in initial plasma aldosterone levels and responses to ACTH, although initial aldosterone levels were significantly higher in rehabilitated seals that died than in seals that lived. Seals with adrenal necrosis associated with herpesvirus infection did not have decreased adrenal hormone responses to ACTH. Differences between initial hormone levels and responses to ACTH in different groups of seals may be associated with differing stress levels. Fecal cortisol assays were not a useful method of assessing adrenal function in these seals, as measured levels did not correlate with plasma cortisol levels.     

INCREASED VIGILANCE TOWARD UNFAMILIAR HUMANS BY HARBOR (PHOCA VITULINA) AND GRAY (HALICHOERUS GRYPUS) SEALS

Two pairs of harbor ( Phoca vitulina ) and three pairs of gray ( Halicboeruls grypus ) seals were exposed to one of three human handlers for 15 min, twice a day, for a total of six sessions. Following habituation to the familiar handler, animals were then exposed to a novel human for 7 min, and then retested for 7 min with the familiar human. In all cases, animals responded to the unfamiliar human with increased vigilant behavior, i. e., they spent more time oriented towards the unfamiliar handler during the first 2 min of the test session than during the same interval of either the final habituation session or the retest with the familiar human ( P = 0.03 in all cases). There was also a tendency for seals to contact the familiar handler with their noses more rapidly than the novel human ( P = 0.06). These results support the hypothesis that phocid seals are capable of discriminating between individual humans in their environment, setting the stage for human-based Pavlovian conditioning.     

HAUL-OUT SELECTION BY PACIFIC HARBOR SEALS (PHOCA VITULINA RICHARDII): ISOLATION AND PERCEIVED PREDATION RISK

The potential for non-aquatic predators to influence habitat use by harbor seals ( Phoca vitulina ) in a nearshore marine environment was studied by examining haul-out site use and through an experimental approach. Distance from shore, distance to possible foraging depths, peripheral water depth, and haul-out areas were quantified for each haul-out. There was a positive relationship between the number of seals hauled out and the distance from shore for eight known haul-out sites. The hypothesis that harbor seals increasingly hauled out farther offshore to reduce predation risk was tested experimentally by measuring their response to a model of a potential terrestrial predator in comparison to a control object, and to disturbance by a human at one of the study sites. Harbor seals abandoned the haul-out in the presence of the predator model, but showed little response to the controls, suggesting they possess a threat image for terrestrial predators and avoid hauling out when it is perceived. These results support the hypothesis that harbor seals select isolated sites to reduce exposure to terrestrial carnivores.     

PLASMA PROGESTERONE CONCENTRATIONS MEASURED USING AN ENZYME-LINKED IMMUNOSORBENT ASSAY USEFUL FOR DIAGNOSING PREGNANCY IN HARBOR SEALS (PHOCA VITULINA)

Wild-caught female harbor seals ( Phoca vitulina ) were classified as sexually mature or immature on the basis of standard body length (< 125 cm immature, > 125 cm mature) and plasma progesterone concentrations measured using an enzyme-linked immunosorbent assay (ELISA), a technique usable in the field. Sexually mature females were classified as pregnant or non-pregnant on the basis of their plasma progesterone concentrations. Of 28 wild mature female harbor seals caught in the Moray Firth, N.E. Scotland, between the end of February and the end of May, 79% had plasma progesterone concentrations greater than 60 nmol liter⁻¹, the lowest plasma progesterone concentration measured in one of eight females later observed with a pup, and were diagnosed as pregnant. A linear discriminant function, calculated to provide a method of distinguishing pregnant and non-pregnant females, predicted 100% of non-pregnant females and 95.8% of pregnant females using plasma progesterone concentration, standard length, and month of capture as parameters. Plasma progesterone concentrations were less than 30 nmol liter⁻¹ in all mature and immature males and immature females. In mature females plasma progesterone concentrations ranged from 0-318 nmol liter⁻¹.     

STABILITY IN THE PROPORTION OF HARBOR SEALS HAULED OUT UNDER LOCALLY IDEAL CONDITIONS

We monitored the haul-out behavior of 68 radio-tagged harbor seals ( Phoca vitulina ) during the molt season at two Alaskan haul-out sites (Grand Island, August-September 1994; Nanvak Bay, August-September 2000). For each site, we created a statistical model of the proportion of seals hauled out as a function of date, time of day, tide, and weather covariates. Using these models, we identified the conditions that would result in the greatest proportion of seals hauled out. Although those "ideal conditions" differed between sites, the proportion of seals predicted to be hauled out under those conditions was very similar (81.3% for Grand Island and 85.7% for Nanvak Bay). The similar estimates for both sites suggest that haul-out proportions under locally ideal conditions may be constant between years and geographic regions, at least during the molt season.     

MOLTING PHENOLOGY OF HARBOR SEALS ON TUGIDAK ISLAND, ALASKA

We documented the progression and timing of the annual molt of harbor seals on Tugidak Island, Alaska, from 1997 to 1999. In all years the timing of molting differed among age-sex classes. Yearlings molted first, subadults second, adult females third, and lastly adult males. Timing of molting was nearly identical in 1997–1998, whereas in 1999 molting occurred three to six days later for all age-sex classes except yearlings. Estimated dates when peak proportions of each age-sex class were molting ranged from 2 August (yearlings) to 2 September (adult males). The number of seals hauled out was positively related to the proportion of seals in the molt and negatively related to the proportion of seals in the postmolt. Population trend estimates, based on aerial counts conducted during a narrow window within the molting period, are likely biased toward certain age-sex classes. Statistical models used to estimate trend include covariates to help account for within-year variation in seal numbers, but do not account for compositional changes that occur during molting. Population modeling may elucidate the effects of within-year population structure on trend estimates. Monitoring molting phenology at additional sites is necessary to determine the extent of geographic variation in molting.     

THE DISTRIBUTION, ABUNDANCE AND SELECTED PREY OF THE HARBOR SEAL, PHOCA VITULINA CONCOLOR, IN SOUTHERN NEW ENGLAND

The seasonal distribution and abundance of harbor seals occurring south of Maine were documented by counting the number of seals at traditional haulout locations. The average number of seals counted during each survey in Massachusetts and New Hampshire was 3,560 ± 255 (95% CI), 1983–1987. The maximum number of seals counted on any individual survey was 4,736 individuals. Fifty percent of all the surveys since January 1985 have resulted in counts greater than 4,000 seals reflecting a 27% increase in the abundance of seals in our study area since that date. Seventy-five percent of the seals in southern New England are located at haulout sites on Cape Cod and Nantucket Island. The largest aggregation of seals in the eastern United States occurs mid-winter at Monomoy Island and adjacent shoals. A single high count of 1,672 seals occurred at this site during the study period. An additional 271–374 seals were also counted in Rhode Island, Connecticut and eastern Long Island Sound during surveys conducted in March 1986 and 1987. The American sandlance Ammodytes americanus was the single dominant prey item of harbor seals in waters adjacent to Cape Cod based on the modified frequency of occurrence of each prey species in scat samples collected from three haulout sites on Cape Cod between 1984–1987. During January and February sandlance was the near exclusive prey item at Monomoy (99%, n= 80). During March and April, the frequency of Atlantic herring Clupea harengus increased in the scat samples at this site. Regional differences in the diet of seals reflect distinct prey communities throughout the study area. Since 1986, the percent occurrence and importance of sandlance in the diet of seals has decreased, reflecting an overall decrease in abundance of this prey species in waters adjacent to Cape Cod. In spite of fluctuations in abundance, and regional differences in the diet of seals throughout the study area, sandlance still comprised a minimum 55% of the total prey species of harbor seals throughout the study area.     

THE MAXIMUM RANGE OF VOCAL COMMUNICATION IN AIR BETWEEN A HARBOR SEAL (PHOCA VITULINA) PUP AND ITS MOTHER

Abstract: The transmission losses with distance of four pure tones (0.5, 1, 2, 4 kHz) were measured in the air 9 cm above the surface of the coastal waters in the Bay of Fundy. The study was conducted between May and July 1992, on warm days with low winds. The measured transmission losses were as much as 11 dB less than predicted by spherical spreading (6 dB/distance doubled), at 400 m (0.5 kHz). This enhanced sound transmission is probably due to the air temperature profile which increases with height above the water's surface. Such a profile causes sound waves to refract towards the water, thereby reducing spreading losses. High-frequency sound absorption negates enhanced transmission at 4 kHz, at distances over 500 m. On days with low winds and low ambient noise levels, a seal pup calling at 90 dB re 20, μPa at 0.5 kHz should be detectable by the mother up to 1 km away, and may be audibly recognizable up to 140 m away.     

CHANGES IN HEART RATE VARIABILITY DURING DIVING IN YOUNG HARBOR SEALS, PHOCA VITULINA

We studied changes in the patterns of heart rate variability (HRV) that coincide with the development of diving skills in harbor seal pups, Phoca vitulina . Heart rate measurements were collected remotely. Spectral analysis of HRV revealed power within a mid-frequency band (0.1–0.3 Hz) which was prominent, especially in pups less than 10 d of age. In these younger pups, the heart rate switched cyclically between a low and a high diving heart rate every 3–10 sec. Older pups exhibited a highly controlled diving bradycardia with a lower median and a lower variance when compared to younger individuals. These results provide new insight into the maturation of the bradycardia component of the dive response in harbor seal pups.     

AGONISTIC BEHAVIOR IN HARBOR SEALS (PHOCA VITULINA) IN RELATION TO THE AVAILABILITY OF HAUL-OUT SPACE

The hypothesis that haul-out space is a contested resource among harbor seals (Phoca vitulina) was tested by comparing the number of agonistic interactions and the time devoted to them, at a haul-out with unlimited space and a haul-out with limited space, in Humboldt County, California. 798 agonistic interactions were recorded in 160 h of observation. The average number of agonistic interactions per seal per hour and the time spent on these interactions were significantly higher at the site with limited haul-our space (P < 0.05, n= 20). At this restricted site, the frequency of agonistic interactions was positively correlated with seal density (r= 0.61, P < 0.05, n= 20), as was time spent on agonistic interactions (r= 0.84, P < 0.05, n= 20). Size, sex, and age-class of individuals all influenced agonistic interactions. In 72.5% of displacements, smaller seals were displaced by larger ones. This may support claims that some harbor seal populations maintain a size-based dominance hierarchy. The results of this study support the hypothesis that there is competition among harbor seals for suitable haul-out space where this resource is limited. The outcome of these interactions may be predicted, at least in part, by theoretical models on pay-off asymmetries, resource holding power, and the prior residency effect.     

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.     

GROWTH AND DEVELOPMENT IN FREE-RANGING HARBOR SEAL (PHOCA VITULINA) PUPS FROM SOUTHERN BRITISH COLUMBIA, CANADA

Harbor seals ( Phoca vitulina ) are small pinnipeds that are widely distributed throughout the temperate coastal regions of the Atlantic and Pacific oceans. We determined birth mass, neonatal growth rates, weaning age, and weaning mass of NE Pacific harbor seals ( P. v. richardsi ) during a capture-recapture study that spanned the nursing period (Sidney Island, British Columbia, Canada). Of 46 harbor seal pups initially captured, 28 were classified as newborns ( i. e. , < 24 h old). Mean body mass of newborns was 11.2 ± SE 0.31 kg. Pups were individually tagged and recaptured throughout the nursing period. Average daily mass gain during the nursing period was 394 ± 26 g. Mean birth mass of males did not differ significantly from females, although pups found with fetal pelage (lanugo) (21.4% of all newborns) were smaller at birth (9.8 ± 0.44 kg) than non-lanugo pups (11.6 ± 0.33 kg). Mean weaning mass was estimated at 23.6 ± 1.2 kg at a mean weaning age of 32 d ± 1.5 d. While birth and weaning masses differed little from the published data for offshore Sable Island harbor seals ( P. v. concolor ), British Columbia harbor seals are characterized by half the daily mass gain and a longer nursing period.     

SUMMER HOME RANGE AND HABITAT SELECTION OF HARBOR SEAL (PHOCA VITULINA) PUPS

We studied the summer home range and habitat selection of harbor seal pups tracked using VHF radio telemetry along the coast of central Norway in 1997 and 1998. Median fixed kernel home range of six pups tracked in 1998 was 10.4 km² and the median core area was 1.2 km². One particular deep basin (>100 m deep) was highly selected, and extensively used by four of the pups, probably for feeding. An area adjacent to the deep basin also was selected, probably for haul outs. One pup spent most of the time in inshore, kelp forest areas. Shallow areas without kelp forest and exposed land were used least frequently, possibly because they did not provide suitable areas for feeding and to haul out. The seals hauled out more during calm wind and low tide than during rough wind and high tide. Seals seemed to haul out on intertidal rocks associated with islands, and alternated between several haul-out sites. We found no pattern in haul-out activity associated with age, time of day, air/sea temperature, cloud cover, or rainfall. In 1997, seven pups were tracked manually. These data were associated with methodological limitations, and they were used only to illustrate that these seals stayed mainly within the shallow, inshore, kelp forest area, indicating a possible difference in habitat use between years.     

DO PHOTOPERIOD AND TEMPERATURE INFLUENCE THE MOLT CYCLE OF PHOCA VITULINA IN CAPTIVITY?

Photoperiod variations are the principal environmental signals entraining circannual activity in mammals, which is also believed to be influenced by changes in temperature and nutrition. Control of the molt in phocids has not been extensively studied with experimental manipulations of environmental factors, but substantial work indicates that photoperiod has a primary influence on mammalian breeding and furring mechanisms. Studies were under taken to reestablish the molt cycle in five harbor seals ( Phoca vitulina ) previously kept in an enclosed facility with a limited air/water temperature gradient and artificial photoperiod. The seals were exposed to an extended temperature gradient during the first period of study, and no molting was observed. The second period of study was characterized by a natural occurring photoperiod of 8 h 53 min light (L)/15 h 07 min dark (D) and a maximal 15 h 31 min L/8h 29 min D duration, and molting occurred in all individuals. These results suggest a prevalent influence of photoperiod variation on the harbor seal molt.