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.     

Efficacy of voluntary mitigation in reducing harbor seal disturbance

Marine and coastal tourism has rapidly expanded worldwide in the past 2 decades, often occurring in once secluded habitats. In Alaska, tourism near tidewater glaciers has attracted millions of visitors and increased the presence of ships, tour vessels, and coastal development. Although sustainable tourism, resulting from balanced effects on wildlife and client satisfaction, is a goal of most tourism operators, it is not always achieved. Voluntary compliance with viewing guidelines and codes of conduct have been encouraged, but few assessments have the longitudinal scope to evaluate long-term changes in impacts on wildlife and the ability of vessel operators and kayak guides to sustain lower impact operating practices over time. This study assessed vessel and kayak visitation and resulting impacts on harbor seals in the Kenai Fjords National Park, southcentral Alaska. We obtained observations from 2002 to 2011, using remotely controlled video cameras located near Aialik and Pedersen Glaciers in the Kenai Fjords National Park. Overall, disturbance was associated with 5.1% of vessel sightings, 28% of vessel interactions (vessel observed within approx. 300 m of seals), 11.5% of kayak sightings, and 61% of kayak interactions. Results demonstrated that voluntary changes in operations significantly reduced vessel and kayak disturbance of seals by 60–80%. Even with prior establishment of operating guidelines, tour vessel captains were able to further reduce their effect on wildlife with more careful operations. Rapid growth of guided kayak excursions that occurred during this study caused greater disturbance to seals than motorized vessels but guide trainings helped reduce disturbances. Diminished impacts of motor vessels and kayakers persisted across years although effects of kayaks were less consistent than motor vessels, which reflected greater variability in inter-annual spatial use patterns by kayakers. Long-term monitoring, including assessments of wildlife responses to vessel and kayak operations, combined with 2-way communication with vessel operators and guides, enhanced the effectiveness of mitigation and facilitated adaptive adjustments to mitigation protocols over time. © 2013 The Wildlife Society.     

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.     

Interannual differences in postrelease movements of rehabilitated harbor seal pups (Phoca vitulina richardii) in the Salish Sea

Despite the large number of harbor seals (Phoca vitulina richardii) rehabilitated worldwide, few studies have been conducted on postrelease movement and behavior of rehabilitated harbor seal pups. We compared interannual differences in movements and survival of 24 rehabilitated seal pups released in the Salish Sea in 2010 (n = 10), 2012 (n = 5), 2013 (n = 5), and 2014 (n = 4). We also compared the postrelease movement of these seals to the movement of 10 wild seal pups tracked in the same ecosystem in 2010. Transmission duration, total cumulative distance, and average daily distance varied annually. Maximum linear distance traveled from the release site was similar for the rehabilitated seal groups. Compared to wild seals (n = 10), and consistent with prior studies, rehabilitated pups (n = 24) traveled significantly farther daily and cumulatively than wild weaned pups. Unlike in a prior study in this ecosystem, we found no significant difference between transmission duration in wild and rehabilitated pups.     

Reaction of Harbor Seals to Cruise Ships

Abstract: The largest aggregations of harbor seals (Phoca vitulina) in Alaska, USA, haul out on floating ice in tidewater glacial fjords. Seals use these fjords in peak numbers during the critical periods of pupping, breeding, and molting when visits by tour ships also peak. Documented and suspected declines of harbor seals in fjords with rising vessel traffic underscore the need to better understand possible impacts, particularly in areas where ship visits have risen substantially in the past 2 decades. We examined the interruption of haul-out bouts of harbor seals due to approaching cruise ships in Disenchantment Bay, Alaska. We conducted observations from cruise ships and focused on disturbance of seals as evidenced by seals flushing into the water from the floating ice on which they rested. We investigated rate of flushing in relation to vessel distance, approach angle, group size, and seal type (mother, pup, or other). Using a survival-regression analysis, we found that the risk of disturbing harbor seals increased when ships approached within 500 m; seals approached as close as 100 m were 25 times more likely to enter the water than seals 500 m from a ship. Seals were 4 times more prone to enter the water when ships were approaching directly rather than passing abeam. Seals responded similarly regardless of group size or seal type. Energetic models indicated a potential to disrupt energy balance and cause thermal stress in disturbed pups if they spent >50% of their time in ice-chilled water. Studies at non-glacial sites suggest that pups spend 40–70% of their time in the water. Voluntary guidelines for approaching seals in Alaska recommend that cruise ships approach ≥91 m (100 yards), a distance at which we show 90% of seals would flush into the water. Our findings indicate a need to develop regulations to maintain a 500-m separation between cruise ships and seals in all Alaskan glacial fjords.     

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.     

REBUILDING SEAL STOCKS IN THE KATTEGAT-SKAGERRAK

The harbor seal ( Phoca vitulina ) population in the Kattegat-Skagerrak area has been dwindling for several centuries due to excessive hunting pressure. Corrected hunting statistics during 1890–1976 are used to estimate changes in population size over the past century. After protection was introduced in the 1960s and 1970s the harbor seal population in the area increased at an exponential rate of 0.12 and exceeded 5,000 animals in 1986. The present rate of population growth is used for modelling the influence of fertility and age-specific mortality. It is found that the observed high rate of increase is only realistic if female fertility rate is very high, the range of juvenile mortality rate is 0.33–0.52 and adult mortality is less than 0.15. Commonly cited higher mortality rates are not realistic in the Kattegat-Skagerrak area.     

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.     

Current population genetics of Japanese harbor seals: Two distinct populations found within a small area

We used microsatellite loci to understand the current population genetics of Japanese harbor seals, and found that they were subdivided into two distinct populations, one in Erimo and the other in eastern Hokkaido, despite being only 150 km apart and having no obvious geographical boundaries between them. Strong population subdivision between the two areas may be due to the small number of haul-out sites and population size, and that a single habitat type is used for haul-out sites in Japan. Also, at Erimo, there are numerous rocky reefs at a single haul-out site that stretch up to 1.3 km offshore from land, while 10 haul-out sites in eastern Hokkaido, separated by a maximum of 30 km, are interspersed along 75 km of the coast. The direction in which the rocky reefs stretch away from land or in which the haul-out sites are facing may be limiting the movement of Japanese harbor seals and causing strong population subdivision between Erimo and eastern Hokkaido. No recent genetic bottlenecks were detected, although the seals in Erimo and eastern Hokkaido were reported to have experienced declines in population size over the past few decades.     

Harbor seal pup dispersal and individual morphology,hematology, and contaminant factors affecting survival

Understanding the factors affecting individual harbor seal (Phoca vitulina) survival is essential for determining population level health risks. We estimated postweaning dispersal, and modeled the effects of morphology, hematology, and blubber contaminants on the survival of recently weaned harbor seal pups using a mark recapture framework. We deployed satellite transmitters on apparently healthy pups captured in San Francisco Bay (SFB, n = 19) and Tomales Bay (TB, n = 7), and pups released after rehabilitation that stranded along the central California coast preweaning (n = 21). Dispersal distances were further than previously reported for harbor seal pups (maximum = 802 km) which has implications for understanding risks to this vulnerable age class. We found differences in body condition, serum immunoglobulin and thyroxine (T4) concentrations, white blood cell count, and blubber organohalogen contamination (OH) among the three groups. Overall, increased T4, decreased OH, and increased mass were associated with greater survival probabilities; whereas, among stranded seals, greater mass gain, shorter time in rehabilitation, and admission to rehabilitation earlier in the season were associated with greater survival probabilities. Attention to these latter factors may improve the success of rehabilitation efforts. For wild pups, reduction of legacy contaminants and direct causes of mortality, such as ship strike, may enhance pup survival.     

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.     

Harbor seal use of glacier ice and terrestrial haul‐outs in the Kenai Fjords,Alaska

Harbor seals, Phoca vitulina, use diverse haul‐out substrates including ice calved by tidewater glaciers. Numbers of seals at glacial and terrestrial haul‐outs on the southeastern Kenai Peninsula, Alaska, were assessed using aerial, vessel, and video surveys. Mean annual abundance at glacial and terrestrial haul‐outs differed temporally. From 2004 to 2011, numbers of seals counted during the molt increased 5.4%/yr at glacial haul‐outs and 9%/yr at terrestrial haul‐outs while numbers of pups increased 5.0%/yr at glacial sites and 1.5%/yr at terrestrial sites. Numbers of seals without pups counted during pupping increased 7.96%/yr at glacial sites and 5.1%/yr at terrestrial sites. Results indicate that pupping and molting locations are not equivalent and population monitoring during the molt does not necessarily reflect habitat association of pupping seals. Ratios of pups to total seals counted during pupping and the subsequent molt were used to contrast habitat use. Low proportions of pups at terrestrial haul‐outs, relative to most glacial haul‐outs, indicate an overall preference for pupping in glacial haul‐outs. High proportions of pups at most glacial sites (during pupping and molting) suggest reduced use of tidewater glacier habitats by nonbreeders and molting seals. Results suggest more seals associate with glacial haul‐outs than currently estimated.     

The role of Steller sea lions in a large population decline of harbor seals

We provide the first direct evidence that Steller sea lions will prey on harbor seals. Direct observations of predation on marine mammals at sea are rare, but when observed rates of predation are extrapolated, predation mortality may be found to be significant. From 1992 to 2002, harbor seals in Glacier Bay declined steeply, from 6,200 to 2,500 (∼65%). After documenting that Steller sea lions were preying on seals in Glacier Bay, we investigated increased predation by sea lions as a potential explanation for the large decline. In five independent data sets spanning 21–25 yr and including 14,308 d of observations, 13 predation events were recorded. We conducted a fine-scale analysis for an intensively studied haul-out (Spider Island) and a broader analysis of all of Glacier Bay. At Spider Island, estimated predation by sea lions increased and could account for the entirety of annual pup production in 5 of 8 yr since 1995. The predation rate, however, was not proportional to the number of predators. Predation by Steller sea lions is a new source of mortality that contributed to the seal declines; however, life history modeling indicates that it is unlikely that sea lion predation is the sole factor responsible for the large declines.     

Determining a correction factor for aerial surveys of harbor seals in California

Counts of pinnipeds provide a minimal estimate of population size because some unknown proportion of individuals is in the water during surveys. We determined a correction factor (CF) for Pacific harbor seals (Phoca vitulina richardii) by estimating the proportion ashore of 180 seals tagged with flipper‐mounted radio tags throughout California. The mean proportions of tagged individuals ashore during four complete surveys in 2004 were not different between central and northern California (F= 1.85, P= 0.18) or between sexes (F= 0.57, P= 0.45), but a lesser proportion of weaners was ashore than subadults or adults (F= 7.97, P= 0.001), especially in northern California. The CF calculated for the statewide census of harbor seals was 1.65, using transmitters operating during the survey (n= 114). Using a mark‐recapture estimator for tag survival (phi) and the four telemetry surveys the mean CF for central and northern California was 1.54 ± 0.38 (95% CI). A CF for southern California of 2.86 was based on a single survey. Using the mean CF of 1.54 and a statewide count in 2009 we estimated 30,196 (95% CI = 22,745–37,647) harbor seals in California.     

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.     

Postrelease dive ability in rehabilitated harbor seals

The efficacy of seal rehabilitation is examined in a postrelease study of dive ability in harbor seal pups (Phoca vitulina) in the Wash, United Kingdom. Six rehabilitated seals were fitted with Sea Mammal Research Unit (SMRU) Argos Satellite Relay Data Logger tags and their individual dive behavior was monitored for an average of 122 d. The upper 90 percentile edge of dive behavior (dive duration [DD₉₀] and percentage of time at‐sea spent in a dive [PD₉₀]), in 7 d bins, was used as a proxy for physiological dive ability. The results are compared with data from five wild adult harbor seals. There was no statistically significant difference between (1) the mean track duration of rehabilitated seals (126.20 ± 27.48 [SD] d) and adult seals (150.2 ± 24.62 d) (P= 0.108), indicating no evidence that short‐term survival was less in the rehabilitated group; (2) the mean mass‐scaled DD₉₀ of rehabilitated seals (3.95 ± 0.37 min) and adult seals (4.09 ± 0.55 min) (P= 0.632); and (3) the mean PD₉₀ of rehabilitated seals (81.62 ± 1.21%) and adult seals (81.48 ± 3.93%) (P= 0.943). These three results all suggest the success of the rehabilitation program in terms of short‐term survival and dive ability.     

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.     

DIFFERENTIAL MOVEMENTS BY HARBOR SEAL PUPS IN CONTRASTING ALASKA ENVIRONMENTS

Movement patterns of Alaska harbor seal pups were studied using satellite telemetry during 1997–2000. Mean tracking duration was 277.3 d (SD = 105.8) for Tugidak Island pups ( n = 26) and 171.2 d (108.3) for Prince William Sound (PWS) pups ( n = 27). Movements were similar for males and females and were largely restricted to the continental shelf. Multiple return trips of > 75 km from the natal area and up to ∼3 wk duration were most common, followed by movements restricted to <25 km from the natal area; one way movements from the natal site were rare. Distances moved and home range sizes remained relatively stable or increased gradually from July through winter, then decreased markedly through spring. Monthly movements (maximum distance from tagging location, mean distance from haul-outs to at-sea locations, and home range size) were significantly greater for Tugidak vs . PWS pups. Six of seven pups from each region that traveled farthest and were tracked the longest had returned to their tagging site when their last location was recorded, indicating philopatry or limited dispersal during their first year of life. Seal pups exhibited similar movement patterns in the distinct habitats of the two regions, but differed in the spatial extent of their movements.     

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.