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.     

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.     

MAJOR DECLINE IN NUMBER OF HARBOR SEALS, PHOCA VITULINA RICHARDSI, ON TUGIDAK ISLAND, GULF OF ALASKA

Tugidak Island, located in the Gulf of Alaska, was once the site of one of largest local concentrations of harbor seals ( Phoca vitulina richardsi ) in the world. This population, which probably consisted of about 20,500 animals in the mid-1960s declined by about 85% between 1976 and 1988. The population appeared to decline more rapidly during the late 1970s than during the 1980s. Causes for the decline are not apparent. There appear to be both similarities and dissimilarities between this decline and recent declines in abundance of northern fur seals ( Callorhinus ursinus ) and Steller sea lions ( Eumetopias jubatus ) in the Bering Sea and Gulf of Alaska.     

THE ABUNDANCE OF HARBOR SEALS IN THE GULF OF ALASKA

The abundance of harbor seals ( Phoca vitulina richardii ) has declined in recent decades at several Alaska locations. The causes of these declines are unknown, but there is concern about the status of the populations, especially in the Gulf of Alaska. To assess the status of harbor seals in the Gulf of Alaska, we conducted aerial surveys of seals on their haul-out sites in August-September 1996. Many factors influence the propensity of seals to haul out, including tides, weather, time of day, and time of year. Because these "covariates" cannot simultaneously be controlled through survey design, we used a regression model to adjust the counts to an estimate of the number of seals that would have been ashore during a hypothetical survey conducted under ideal conditions for hauling out. The regression, a generalized additive model, not only provided an adjustment for the covariates, but also confirmed the nature and shape of the covariate effects on haul-out behavior. The number of seals hauled out was greatest at the beginning of the surveys (mid-August). There was a broad daily peak from about 1100–1400 local solar time. The greatest numbers were hauled out at low tide on terrestrial sites. Tidal state made little difference in the numbers hauled out on glacial ice, where the area available to seals did not fluctuate with the tide. Adjusting the survey counts to the ideal state for each covariate produced an estimate of 30,035 seals, about 1.8 times the total of the unadjusted counts (16,355 seals). To the adjusted count, we applied a correction factor of 1.198 from a separate study of two haul-out sites elsewhere in Alaska, to produce a total abundance estimate of 35,981 (SE 1,833). This estimate accounts both for the effect of covariates on survey counts and for the proportion of seals that remained in the water even under ideal conditions for hauling out.     

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.     

DISPERSAL and BYCATCH MORTALITY IN GRAY, HALICHOERUS GRYPUS, AND HARBOR, PHOCA VITULINA, SEALS TAGGED AT THE NORWEGIAN COAST

Between 1975 and 1998, 3,571 gray and 630 harbor seal pups were tagged along the Norwegian coast, and 259 (7%) gray and 80 (13%) harbor seal tags were returned. Incidental mortality, mainly in bottom-set nets, accounted for the majority of deaths (79% in gray and 48% in harbor seals, respectively). Seals were most vulnerable to incidental mortality in fishing gear during the first three months after birth, but high incidental mortality prevailed during the first 8–10 mo. Gray seals dispersed more widely (mean distance: 120 km) than harbor seals (mean distance: 69 km). Both species dispersed most widely during the two first months after tagging. The maximum distance moved was 739 km for gray and 463 km for harbor seals. Strong fidelity for their place of birth was observed in adult gray seals during breeding season. No significant difference in incidental mortality was detected between the areas of tagging. However, for 37 harbor seals tagged in a 724 km nature reserve no returns were reported.     

AUTONOMIC CONTROL OF HEART RATE AFTER EXERCISE IN TRAINED WRESTLERS

The objective of this study was to establish differences in vagal reactivation, through heart rate recovery and heart rate variability post exercise, in Brazilian jiu-jitsu wrestlers (BJJW). A total of 18 male athletes were evaluated, ten highly trained (HT) and eight moderately trained (MT), who performed a maximum incremental test. At the end of the exercise, the R-R intervals were recorded during the first minute of recovery. We calculated heart rate recovery (HRR_60s), and performed linear and non-linear (standard deviation of instantaneous beat-to-beat R-R interval variability – SD1) analysis of heart rate variability (HRV), using the tachogram of the first minute of recovery divided into four segments of 15 s each (0-15 s, 15-30 s, 30-45 s, 45-60 s). Between HT and MT individuals, there were statistically significant differences in HRR_60s (p <0.05) and in the non linear analysis of HRV from SD1_30-45s (p <0.05) and SD1_45-60s (p <0.05). The results of this research suggest that heart rate kinetics during the first minute after exercise are related to training level and can be used as an index for autonomic cardiovascular control in BJJW.     

DEVELOPMENT OF DIVING BY HARBOR SEAL PUPS IN TWO REGIONS OF ALASKA: USE OF THE WATER COLUMN

Satellite-linked dive recorders were attached to 53 harbor seal pups in Prince William Sound (PWS) and at Tugidak Island, Alaska, during 1997–1999. We used generalized additive models and bootstrap techniques to describe pup diving behavior during their first year of life. Pups increased their ability to dive during the first 3–6 mo, as indicated by increases in proportion of time in the water (time wet) and maximum dive depth achieved by a pup each day (max-depth) values. Time wet and/or max-depth later decreased, suggesting a seasonal component to diving behavior. Monthly time wet varied from an overall minimum of 0.68 at tagging in July to a maximum of 0.89 in November. Pups spent half of their time wet swimming in water <25 m deep, the shallowest 30% of the available water column. They spent only 5% of their time swimming in the deepest 30% of the available water column, at depths >60–70 m. This strongly suggests they were not feeding on or near bottom during their first year. Average max-depths and deepest actual dives were similar for PWS and Tugidak pups. PWS pups dove deeper sooner and spent less time wet than Tugidak pups during the first few months after tagging, probably as a result of regional bathymetric differences. Diving behavior and body condition suggest that food availability was not likely a major factor in the population decline in PWS during the period of this study.     

DIEL HAUL-OUT PATTERNS AND SITE FIDELITY OF HARBOR SEALS (PHOCA VITULINA RICHARDSI) ON SAN MIGUEL ISLAND, CALIFORNIA, IN AUTUMN

We studied the haul-out patterns and movements of harbor seals ( Phoca vitulina richardsi ) on San Miguel Island, California, from 23 October through 6 December 1982 by attaching a radio transmitter to each of 18 seals and monitoring their presence ashore with continuously scanning receivers. Seals hauled out at all hours although, on average, the largest proportion of tagged seals was ashore between 1300 and 1500 h. Median durations of haul-out bouts of individual seals ranged from 4.7 to 21.8 h; 81% of all haul-out bouts were less than 12 h and 3% were longer than 24 h. Eighty-one percent of the seals that were resighted at least twice used only the sites where they were tagged; two seals used two sites and one seal used three. Most seals were hauled out on fewer than 51% of the days sampled. On average, about 41% of tagged seals hauled out each day whereas an average of about 19% was hauled out during peak afternoon hours. Using telemetry data to correct a count of 412 seals made during an aerial survey, we estimated absolute abundance at about 2,168 seals; a modified Peterson mark-recapture model produced an estimate of about 1,445 seals.     

SEASONAL CHANGES IN BLUBBER DISTRIBUTION IN ATLANTIC HARBOR SEALS: INDICATIONS OF THERMODYNAMIC CONSIDERATIONS

Among its functions, the hypodermal blubber layer of pinnipeds serves as both an energy reserve and insulation. This study examined seasonal changes in blubber distribution and body morphology in a group of captive changesharbor seals to test whether these changes were designed to maximize insulative effectiveness. Seasonal changes were found in girth, blubber volume, mean blubber depth, and the ratio of blubber depth to body radius (d/r ratio). These changes were more evident in older seals. The d/r ratio demonstrated a smaller relative annual change than mean blubber depth. The d/r ratio also exhibited less variation along the length of the seal than blubber depth at any given time. Similar to reports for ringed seals, and contrary to those for southern elephant seals, the harbor seals preferentially lost blubber from overinsulated areas of the body. These results suggest that core tissue and blubber mass are lost in a manner that maximizes insulative effectiveness.     

CHANGES IN ABUNDANCE OF HARBOR SEALS IN MAINE, 1981–2001

Aerial counts of harbor seals ( Pboca vitulina concolor ) on ledges along the Maine coast were conducted during the pupping season in 1981, 1986, 1993, 1997, and 2001. Between 1981 and 2001, the uncorrected counts of seals increased from 10,543 to 38,014, an annual rate of 6.6 percent. In 2001 30 harbor seals were captured and radio-tagged prior to aerial counts. Of these, 19 harbor seals (six adult males, two adult females, seven juvenile males, and four juvenile females) were available during the survey to develop a correction factor for the fraction of seals not observed. The corrected 2001 abundance estimate was 99,340 harbor seals. Productivity in this population has increased since 1981 from 6.4% pups to 24.4% pups. The number of gray seals ( Halichoerus grypus ) counted during the harbor seal surveys increased from zero in both 1981 and 1986 to 1,731 animals in 2001.     

TRENDS IN ABUNDANCE OF ALASKA HARBOR SEALS, 1983–2001

We estimated trends in abundance of harbor seals ( Phoca vitulina richardsii ) using over dispersed, multinomial models and counts obtained during aerial surveys conducted during 1983–2001 in the Ketchikan, Sitka, Kodiak, and Bristol Bay areas of Alaska. Harbor seal numbers increased significantly at 7.4%/yr during 1983–1998 and 5.6%/yr during 1994–1998 in the Ketchikan area, and 6.6%/yr during 1993–2001 in the Kodiak area. Counts were stable (trends not significant) during 1984–2001 (0.7%/yr) and 1995–2001 (-0.4%/yr) in Sitka, and during 1998–2001 (-1.3%/yr) in Bristol Bay. The influence of covariates ( e.g. , survey date, tide height) on trend estimates was significant and varied among areas and across years, demonstrating the need to include covariates in statistical analyses to accurately estimate trend. Our increasing trend estimate for Kodiak represents the first documented increase in harbor seal numbers over a relatively expansive area in the Gulf of Alaska. However, the trend for the Gulf of Alaska stock is equivocal due to the continued decline in Prince William Sound. Similarly, the trend for the Southeast Alaska stock is equivocal based on our increasing (Ketchikan) and stable (Sitka) trend estimates, and a recent decline reported for Glacier Bay. The Bering Sea stock appears stable after a period of possible decline.     

DEVELOPMENT OF DISPLAY BEHAVIOR IN YOUNG CAPTIVE BEARDED SEALS

In this study of the ontogeny of vocal behavior in captive bearded seals, Erignathus barbatus , (three males and three females), only males exhibited vocal displays. The onset of display behavior coincided with sexual maturity. Males exhibited three types of dive displays associated with the performance of vocalizations. Vocalizing individuals were frequently attended by another male that maintained passive muzzle contact with the vocalizing male. These interactions were non-aggressive and might play a role in the establishment of a social hierarchy or they might allow the attendee to obtain "near-field" vocal information from the displaying male. Captive males' vocalizations resembled those of males in the wild. However, display dives were shorter, and fewer vocalization types were documented among the captive males compared to bearded seals in the wild. The capacity of the captive males for producing well-formed, long calls with large frequency changes was also significantly less than for wild males. These capacities will likely develop further as the males grow older. Individual capacity for vocal production appears to develop gradually, showing plasticity in form development over time.     

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.     

MOVEMENTS OF SATELLITE-TAGGED SUBADULT AND ADULT HARBOR SEALS IN PRINCE WILLIAM SOUND, ALASKA

Satellite-linked tags were attached to 49 subadult and adult harbor seals captured in Prince William Sound (PWS), Alaska, and their movements were monitored during 1992–1997. Seals were tracked for a total of 5,517 seal-days and were located on about 80% of the days that tags transmitted. Most locations were in or near PWS, but some juvenile seals moved 300–500 km east and west into the Gulf of Alaska. While several seals travelled to 50–100 km offshore, virtually all locations were in water <200 m deep. Overall, juvenile seals moved more than adults and had larger home ranges. Movements were significantly affected by month, and age by month and sex by month interactions. In all months, mean distances between successively used haulouts were <10 km for adults and <20 km for juveniles. Mean monthly home ranges varied from <100 km² to >1,500 km², and were smallest during June-July. Mean haul-out to at-sea distance was 5–10 km for adults and generally 10–25 km for juveniles. Satellite-linked tags provided an effective means of monitoring and describing the full range of harbor seal movements in this region, with the exception of late summer when tags were shed during the molt.     

不同年龄健康被试者心率变异性的谱分析研究

对不同年龄健康男性被试者卧、立位条件下,心率变异性（ｈｅａｒｔ ｒａｔｅ ｖａｒｉａｂｉｌｉｔｙ,ＨＲＶ）谱高、低频成分随年龄变化的规律进行研究。结果表明：虽然各年龄组间卧、立位时的静息平均心率并无显著差别,但ＨＲＶ却随年龄的增长而明显降低,青年组ＨＲＶ谱高、低频谱峰功率的显著高于中年组、中老年组及老年组（Ｐ〈０．０５或Ｐ〈０．０１）;ＨＲＶ谱低频谱功率、高频谱功率及总功率均随年龄增长呈指数衰减规     

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.     

TRENDS IN ABUNDANCE AND CURRENT STATUS OF HARBOR SEALS IN OREGON: 1977–2003

The distribution and abundance of harbor seals ( Phoca vitulina richardii ) in Oregon were monitored from 1977 to 2003 by aerial photographic surveys. Harbor seals on shore were counted each year during the reproductive period. Mean annual counts of non-pups (adults and subadults) were used as an index of population size and the trend in the counts was modeled using exponential (density-independent) and generalized logistic (density-dependent) growth models. Models were fit using maximum likelihood and evaluated using Akaike's Information Criterion. The population dynamics of harbor seals in Oregon were best described by the generalized logistic model. The population grew following protection under the Marine Mammal Protection Act of 1972 until stabilizing in the early 1990s. The estimated absolute abundance of harbor seals (all age classes) during the 2002 reproductive period was 10,087 individuals (95% confidence interval was 8,445–12,046 individuals). The current predicted population size for harbor seals in Oregon is above its estimated maximum net productivity level and hence within its optimum sustainable population range. We speculate that recent increases in ocean productivity in the eastern Pacific Ocean may lead to an increase in carrying capacity and renewed growth in Oregon's harbor seal population.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

CORRECTING AERIAL SURVEY COUNTS OF HARBOR SEALS (PHOCA VITULINA RICHARDSI) IN WASHINGTON AND OREGON

Aerial surveys of harbor seals on land produce only a minimum assessment of the population; a correction factor to account for the missing animals is necessary to estimate total abundance. In 1991 and 1992, VHF radio tags were deployed on harbor seals ( n = 124) at six sites in Washington and Oregon. During aerial surveys a correction factor to account for seals in the water was determined from the proportion of radio-tagged seals on shore during the pupping season. This proportion ranged from 0.54 to 0.74. Among the six sites there was no significant difference in the proportion of animals on shore nor was there a difference in age/sex categories of seals on shore between sites. The pooled correction factor for determining total population abundance was 1.53. An additional 32 seals were radio tagged in 1993 at one of the sites used in 1991. Comparing data from the two years, we found no interannual variation. Aerial surveys of all known harbor seal haul-out sites in Washington ( n = 319) and Oregon ( n = 68) were flown during the peak of the pupping season, 1991–1993. The Washington and Oregon harbor seal population was divided into two stocks based on pupping phenology, morphometics, and genetics. Mean counts for the Washington inland stock were 8,710 in 1991, 9,018 in 1992, and 10,092 in 1993. Oregon and Washington coastal stock mean counts were 18,363 in 1991, 18,556 in 1992, and 17,762 in 1993. Multiplying the annual count by the correction factor yielded estimates of harbor seal abundance in the Washington inland stock of 13,326 (95% CI = 11,637–15,259) for 1991, 13,798 (95% CI = 11,980–15,890) for 1992, and 15,440 (95% CI = 13,382–17,814) for 1993. In the Oregon and Washington coastal stock the corrected estimate of harbor seal abundance was 28,094 (95% CI = 24,697–31,960) in 1991, 28,391 (95% CI = 24,847–32,440) for 1992, and 27,175 (95% CI = 23,879–30,926) for 1993.