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.     

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.     

ABUNDANCE OF RINGED SEALS (PUSA HISPIDA) IN THE FJORDS OF SPITSBERGEN, SVALBARD, DURING THE PEAK MOLTING PERIOD

Ringed seal (Pusa hispida) abundance in Spitsbergen, Svalbard, was estimated during the peak molting period via aerial, digital photographic surveys. A total of 9,145 images, covering 41.7%–100% of the total fast‐ice cover (1,496 km²) of 18 different fjords and bays, were inspected for the presence of ringed seals. A total of 1,708 seals were counted, and when accounting for ice areas that were not covered by images, a total of 3,254 (95% CI: 3,071–3,449) ringed seals were estimated to be hauled out during the surveys. Extensive behavioral data from radio‐tagged ringed seals (collected in a companion study) from one of the highest density fjords during the molting period were used to create a model that predicts the proportion of seals hauled out on any given date, time of day, and under various meteorological conditions. Applying this model to the count data from each fjord, we estimated that a total of 7,585 (95% CI: 6,332–9,085) ringed seals were present in the surveyed area during the peak molting period. Data on interannual variability in ringed seal abundance suggested higher numbers of seals in Van Keulenfjorden in 2002 compared to 2003, while other fjords with very stable ice cover showed no statistical differences. Poor ice conditions in general in 2002 probably resulted in seals from a wide area coming to Van Keulenfjorden (a large fjord with stable ice in 2002). The total estimated number of ringed seals present in the study area at the time of the survey must be regarded as a population index, or at least a minimum estimate for the area, because it does not account for individuals leaving and arriving, which might account for a considerable number of animals. The same situation is likely the case for many other studies reporting aerial census data for ringed seals. To achieve accurate estimates of population sizes from aerial surveys, more extensive knowledge of ringed seal behavior will be required.     

An assessment of population size and distribution of harbour seals in the Republic of Ireland during the moult season in August 2003

The status of Ireland's harbour seal Phoca vitulina vitulina population is poorly understood. The most recent national population estimate dates back to the breeding season in 1978 and did not cover the entire coastline. Reliable up-to-date information on the abundance and distribution of harbour seals in Ireland is necessary to assess the conservation status of the species and for the effective identification, management and monitoring of special areas of conservation required for harbour seals under the EU Habitats Directive. To provide comprehensive current information on Ireland's harbour seal population, a geographically extensive survey was conducted along the coastline of the Republic of Ireland during the species' annual moult in August 2003. This complemented a similar survey of Northern Ireland, which was conducted in 2002. Using thermal imaging technology, haul-out groups of harbour seals and grey seals Halichoerus grypus were identified from the air, aerial-counts were obtained and compared with simultaneous ground-count data from selected sites. Harbour seal distribution recorded during the 2003 moult season was concentrated in the south-west, west and north-west of the country. This national survey yielded a minimum population estimate for the Republic of Ireland of 2905 harbour seals, delivering an effective baseline for current and future population research.     

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.     

Harbor seal population decline in the Aleutian Archipelago

Populations of Steller sea lions, northern fur seals, and northern sea otters declined substantially during recent decades in the Bering Sea and Aleutian Islands region, yet the population status of harbor seals has not been assessed adequately. We determined that counts obtained during skiff‐based surveys conducted in 1977–1982 represent the earliest estimate of harbor seal abundance throughout the Aleutian Islands. By comparing counts from 106 islands surveyed in 1977–1982 (8,601 seals) with counts from the same islands during a 1999 aerial survey (2,859 seals), we observed a 67% decline over the ～20‐yr period. Regionally, the largest decline of 86% was in the western Aleutians (n= 7 islands), followed by 66% in the central Aleutians (n= 64 islands), and 45% in the eastern Aleutians (n= 35 islands). Harbor seal counts decreased at the majority of islands in each region, the number of islands with >100 seals decreased ～70%, and the number of islands with no seals counted increased ～80%, indicating that harbor seal abundance throughout the Aleutian Islands was substantially lower in the late 1990s than in the 1970s and 1980s.     

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.     

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.     

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.     

Responses of two marine top predators to an offshore wind farm

Quantifying the likely effects of offshore wind farms on wildlife is fundamental before permission for development can be granted by any Determining Authority. The effects on marine top predators from displacement from important habitat are key concerns during offshore wind farm construction and operation. In this respect, we present evidence for no significant displacement from a UK offshore wind farm for two broadly distributed species of conservation concern: common guillemot (Uria aalge) and harbor porpoise (Phocoena phocoena). Data were collected during boat‐based line transect surveys across a 360 km² study area that included the Robin Rigg offshore wind farm. Surveys were conducted over 10 years across the preconstruction, construction, and operational phases of the development. Changes in guillemot and harbor porpoise abundance and distribution in response to offshore wind farm construction and operation were estimated using generalized mixed models to test for evidence of displacement. Both common guillemot and harbor porpoise were present across the Robin Rigg study area throughout all three development phases. There was a significant reduction in relative harbor porpoise abundance both within and surrounding the Robin Rigg offshore wind farm during construction, but no significant difference was detected between the preconstruction and operational phases. Relative common guillemot abundance remained similar within the Robin Rigg offshore wind farm across all development phases. Offshore wind farms have the potential to negatively affect wildlife, but further evidence regarding the magnitude of effect is needed. The empirical data presented here for two marine top predators provide a valuable addition to the evidence base, allowing future decision making to be improved by reducing the uncertainty of displacement effects and increasing the accuracy of impact assessments.     

Maximum likelihood population size estimation of harbour seals in the Dutch Wadden Sea based on a mark–recapture experiment

1. The harbour seal population Phoca vitulina in the entire Wadden Sea was severely depleted due to a virus-epizootic during 1988. A comprehensive study on the population biology and activity patterns was subsequently initiated to design a management and conservation plan. The main objective of this study was to estimate harbour seal abundance in the different regions of the Wadden Sea.
2. We investigated the potential of a mark–recapture experiment using VHF radio-tags in combination with repeated aerial surveys to estimate the number of harbour seals in the Dutch part of the Wadden Sea. The number of harbour seals hauled-out and the presence of any radio-tagged seals was monitored during seven aerial surveys of all known haul-out sites in the Dutch Wadden Sea over the 1994 breeding season.
3. A maximum likelihood (ML) estimator was developed to infer the rate of tag-loss and the size of the local prepupping population.
4. The ML estimate of the number of harbour seals in the Dutch Wadden Sea was 1536 (95% confidence limits were 1225 and 2200). The corresponding maximum proportion of seals hauled-out was 68%.
5. The use of VHF radio-tags which can be monitored from the air provides a way of correcting aerial survey counts for the proportion of harbour seals hauled-out during the surveys. Since haul-out behaviour may be influenced by local conditions, such as exposure time of sand banks, we recommend this technique be repeated in other areas of the Wadden Sea rather than using the estimates from the current study in other areas.     

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.     

Haul-out behavior of harbor seals (Phoca vitulina) in Hood Canal, Washington

The goal of this study was to model haul-out behavior of harbor seals (Phoca vitulina) in the Hood Canal region of Washington State with respect to changes in physiological, environmental, and temporal covariates. Previous research has provided a solid understanding of seal haul-out behavior. Here, we expand on that work using a generalized linear mixed model (GLMM) with temporal autocorrelation and a large dataset. Our dataset included behavioral haul-out records from archival and VHF radio tag deployments on 25 individual seals representing 61,430 seal hours. A novel application for increased computational efficiency allowed us to examine this large dataset with a GLMM that appropriately accounts for temporal autocorellation. We found significant relationships with the covariates hour of day, day of year, minutes from high tide and year. Additionally, there was a significant effect of the interaction term hour of day : day of year. This interaction term demonstrated that seals are more likely to haul out during nighttime hours in August and September, but then switch to predominantly daylight haul-out patterns in October and November. We attribute this change in behavior to an effect of human disturbance levels. This study also examined a unique ecological event to determine the role of increased killer whale (Orcinus orca) predation on haul-out behavior. In 2003 and 2005 these harbor seals were exposed to unprecedented levels of killer whale predation and results show an overall increase in haul-out probability after exposure to killer whales. The outcome of this study will be integral to understanding any changes in population abundance as a result of increased killer whale predation.     

Estimation of In-Water Density and Abundance of Harbor Seals

Ecologists and managers require accurate population estimates of marine mammals to assess potential anthropogenic threats to these animals. We present estimates of in-water density and abundance of a distinct stock of harbor seals (Phoca vitulina richardii) in Hood Canal, Washington, USA. We used aerial line-transect survey data collected from 2013 to 2016 to directly estimate harbor seal density and abundance in the waters of Hood Canal, a deep-water fjord in the Salish Sea. We estimated a correction factor for trackline detection probability from dive and surface time data gathered from regional seal tagging studies, and applied this factor to correct for seals missed on the trackline during surveys. We applied conventional and multiple covariate line-transect approaches in the analysis. The resulting best estimate of in-water density of harbor seals in the Hood Canal study region was 5.80 seals/km², with an estimated abundance of 2,009 seals. We did not derive a correction factor to account for the number of seals on land (i.e., hauled out). Therefore, these estimates do not reflect total stock size but provide a starting point to evaluate potential influences of anthropogenic activities, particularly those involving underwater noise, on this marine mammal stock. © 2021 The Wildlife Society.     

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.     

EFFECTS OF AN OFFSHORE OIL DEVELOPMENT ON LOCAL ABUNDANCE AND DISTRIBUTION OF RINGED SEALS (PHOCA HISPIDA) OF THE ALASKAN BEAUFORT SEA1

This study investigates how densities of ringed seals were affected by construction and oil production activities at Northstar, an artificial island built in the nearshore Alaskan Beaufort Sea. Intensive and replicated aerial surveys of seals on landfast ice were conducted during six spring seasons: for three seasons before island construction began (1997–1999); after a winter of intensive island construction (2000); and after more limited construction plus drilling (2001) and drilling plus oil production (2002). A Poisson regression model was used to examine seal densities relative to distance from Northstar after allowance for environmental covariates. Post hoc power analysis indicated that the study design and Poisson regression approach had high power to detect small‐scale changes in seal densities near Northstar if such changes had occurred. However, seal densities during spring were not significantly affected by proximity to Northstar in 2000–2002. Habitat, temporal, and weather factors did have significant effects on seal densities. This study shows that effects of the Northstar oil development on local distribution of basking ringed seals are no more than slight, and are small relative to the effects of natural environmental factors. An understanding of environmental effects is essential when assessing potential impacts of industrial activity on ringed seals.     

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.     

HARBOR SEAL TRACKING AND TELEMETRY BY SATELLITE

We tested a satellite Platform Transmitter Terminal (PTT) in the laboratory (on a float and on captive seals) and on a free-ranging harbor seal in the Southern California Bight to investigate the utility of satellite telemetry in documenting seals'at-sea behavior and movements. We used records from a microprocessor-based time-depth recorder (TDR) to interpret location and diving records from the PTT. For the free-ranging harbor seal, we obtained at least one uplink during 70% (while the seal was at sea) to 82% (while she was ashore) of satellite passes and at least one location each day. Of 62 locations determined by Service Argos for the free-ranging seal, 20 were verified from TDR records to have been at sea; these indicated that the seal may have ranged up to 48 km from the haul-out site, although most locations were within 5 km. The accuracies of locations calculated when the seal was at sea (±15 km) were substantially less than when it was ashore (±1.5 km), thus limiting at-sea tracking of seals by satellite to rather gross movements. Fewer transmissions were detected and locations calculated when the seal was actively diving than when it was swimming near the surface as it departed from or returned to the haul-out site. Consequently, average dive durations indicated by the PTT were substantially shorter than those calculated from TDR records. Documentation of foraging areas and detailed at-sea movements using satellite technology may not be possible for pinnipeds unless PTT-transmission rates are increased substantially from the 1 per 45 set maximum rate now permitted by Service Argos.     

MONITORING THE TREND OF HARBOR SEALS IN PRINCE WILLIAM SOUND,ALASKA, AFTER THE EXXON VALDEZ OIL SPILL

We used aerial counts to monitor the trend in numbers of harbor seals, Phoca vitulina richardsi, in Prince William Sound, Alaska, following the 1989 Exxon Valdez oil spill. Repetitive counts were made at 25 haul-out sites during the annual molt period each year from 1990 through 1997. A generalized linear model indicated that time of day, date, and time relative to low tide significantly affected seal counts. When Poisson regression was used to adjust counts to a standardized set of survey conditions, results showed a highly significant decline of 4.6% per year. Unadjusted counts indicated a slight, but not statistically significant, decline in the number of seals. The number of harbor seals on the trend-count route in eastern and central PWS has been declining since at least 1984, with an overall population reduction of 63% through 1997. Programs to monitor long-term changes in animal population sizes should account for factors that can cause short-term variations in indices of abundance. The inclusion of such factors as covariates in models can improve the accuracy of monitoring programs.     

THE MOST NORTHERLY HARBOR SEAL, PHOCA VITULINA, AT PRINS KARLS FORLAND, SVALBARD

The harbor seal ( Phoca vitulina ) has its northernmost distribution at the Norwegian arctic archipelago of Svalbard. Little information exists on this particular harbor seal population. The present paper summarizes this information, and gives the result of surveys of harbor seals conducted in Svalbard in 1984, 1985 and 1987. These surveys show that harbor seals in Svalbard are limited to the area around Prins Karls Forland, the westernmost island in the archipelago. The harbor seal population at Prins Karls Forland numbers at least five to six hundred animals.