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.     

Dive types and circadian behaviour patterns of Saimaa ringed seals<Emphasis Type="Italic">Phoca hispida saimensis</Emphasis> during the open-water season

Diving and circadian behaviour patterns of 7 free-ranging Saimaa ringed sealsPhoca hispida saimensis Nordquist, 1899 were examined by VHF-radiotelemetry during open-water seasons between May and November in Lake Saimaa, eastern Finland. The mean recorded dive duration ranged from 2.8 to 6.5 min, with a maximum of 21 min. The mean dive depth ranged from 9.8 to 15.7 m, with maximum of 39.6 m. The maximum dive depth of each seal was limited by water depth in the study area. The dive depths were positively correlated with dive duration and body mass of the seal. Five different dive types were defined, as based on their depth-time characteristics, each falling into one of the three functional categories: travelling, feeding, and resting. Long duration diving bouts occurred mostly at night and were presumed to be resting dives. Saimaa ringed seals exhibited a circadian pattern of haul-out behaviour that shifted seasonally. During molting (May–June) the seals hauled-out both day and night, but later in summer haul-out was more frequent at night.     

Harbour seal grouping as an anti-predator strategy

Terrestrial grouping by western Atlantic harbour seals, Phoca vitulina concolor, was investigated to determine whether it has a functional significance and, if so, what the function might be. Two nofunctional hypotheses were tested: (1) groups arise from a shortage of haul-out sites or space on sites; and (2) terrestrial groups are extensions of groups formed in the water for reasons not related to hauling-out. Aerial surveys of haul-out site use by seals indicated that sites were not in short supply. Observations of the use of space on one haul-out site suggested that space on sites was not limiting. Arrivals and departures of seals at one study site showed that groups formed at the haul-out site and disbanded after the haul-out period. The hypothesis that grouping functions to increase the probability of predator detection was tested by measuring individual and group vigilance relative to group size, and simulating predator attacks in the field. Although individual vigilance decreased with increasing group size, group vigilance increased. Larger groups detected ‘attacks’ at greater distances and a solitary seal failed to detect an ‘attack’. It is proposed that an individual increases its probability of detecting a predator by joining a group.     

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.     

Fine-Scale Variability in Harbor Seal Foraging Behavior

Understanding the variability of foraging behavior within a population of predators is important for determining their role in the ecosystem and how they may respond to future ecosystem changes. However, such variability has seldom been studied in harbor seals on a fine spatial scale (<30 km). We used a combination of standard and Bayesian generalized linear mixed models to explore how environmental variables influenced the dive behavior of harbor seals. Time-depth recorders were deployed on harbor seals from two haul-out sites in the Salish Sea in 2007 (n = 18) and 2008 (n = 11). Three behavioral bout types were classified from six dive types within each bout; however, one of these bout types was related to haul-out activity and was excluded from analyses. Deep foraging bouts (Type I) were the predominant type used throughout the study; however, variation in the use of bout types was observed relative to haul-out site, season, sex, and light (day/night). The proportional use of Type I and Type II (shallow foraging/traveling) bouts differed dramatically between haul-out sites, seasons, sexes, and whether it was day or night; individual variability between seals also contributed to the observed differences. We hypothesize that this variation in dive behavior was related to habitat or prey specialization by seals from different haul-out sites, or individual variability between seals in the study area. The results highlight the potential influence of habitat and specialization on the foraging behavior of harbor seals, and may help explain the variability in diet that is observed between different haul-out site groups in this population.     

Daily and seasonal variation in the haul-out behavior of the South American sea lion

In the South American sea lion (Otaria flavescens), daily fluctuations in abundance have been analyzed based on sequential counts of the number of animals hauled-out. However, no studies have analyzed haul-out activity in relation to an annual cycle or according to different age/sex classes. The objective of this study was to determine the daily and seasonal haul-out patterns of each age/sex class of South American sea lions as a function of the time of the day. A total of 222 days were analyzed in a breeding colony of Chile, from May 2008 to December 2010. During the non-breeding season (March to December) males, females, and juveniles showed a unimodal pattern, in which few sea lions are hauled-out in the morning and maximum numbers are found in the rookery during the early afternoon (1330–1630). In contrast, during the breeding season (austral summer) the proportion of individuals hauled-out shifted from a unimodal to a bimodal pattern, especially in the case of juveniles. Our results indicate that there are fine scale differences in haul-out behavior among age/sex classes, as well as larger scale seasonal differences in the proportion of sea lions ashore. These differences appear to be related to reproductive activities, food availability and thermoregulatory requirements. These patterns of seasonal variability of South American sea lion haul-out should be taken into consideration when planning surveys to estimate population abundance.     

The effect of a large Danish offshore wind farm on harbor and gray seal haul-out behavior

This study investigates the effects of the construction and operation of a large Danish offshore wind farm on harbor and gray seal haul-out behavior within a nearby (4 km) seal sanctuary. Time-lapse photography, visual monitoring, and aerial surveys were used to monitor the number of seals on land in daylight hours. Seals were monitored during two preconstruction periods (19 June–31 August 2001 and April–August 2002), a construction period of the wind farm (August 2002–December 2003), and a period of operation of the wind farm (December 2003–December 2004). Monthly aerial surveys were conducted to estimate the proportion of seals in the sanctuary relative to neighboring haul-out sites. From preconstruction to construction and through the first year of operation the number of harbor seals in the sanctuary increased at the same rate as the number of seals at the neighboring haul-out sites. No long-term effects on haul-out behavior were found due to construction and operation of the wind farm. However, a significant short-term decrease was seen in the number of seals present on land during sheet pile driving in or near the wind farm. Acoustic deterrents were utilized simultaneously to avoid hearing damage.     

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.     

Movements, diving and foraging behaviour of grey seals (Halichoerus grypus)

This study presents the first direct observations of the movements and behaviour of free-ranging grey seals at sea. Radio and ultrasonic transmitters were attached to three sub-adult male grey seals which were then tracked from a suitable vessel. Behaviour at sea fell into one of three categories: travelling between haul-outs, short duration trips, and resting adjacent to haul-out sites. Travelling was characterized by direct, relatively fast horizontal movement and by V-shaped dive profiles. During short duration trips the seals swam slower and invariably exhibited square-wave dive profiles, spending approximately 60% of total dive duration at the maximum depth. Resting involved shallow dives close to haul-out sites and an absence of directed lateral movement.
The excellent navigational abilities of grey seals are illustrated by the rapid, direct swimming between distant haul-out sites. It is proposed that short duration trips are specifically for foraging because of their association with other piscivores, and because swimming was slow and mostly on or near the sea bed (grey seals are known to feed almost exclusively on demersal and benthic fish). These trips accounted for only 14% of the nine days that seal 1 was tracked. It is also proposed that the habit of diving to the sea bed whilst travelling between distant haul-out sites is to allow opportunistic foraging with only a small increase in total swimming distance.     

Influence of time of day and month on Weddell seal haul-out patterns at the Vestfold Hills, Antarctica

A diurnal pattern in haul-out behaviour has been described for Weddell seals at McMurdo Sound, Antarctica, but regional and seasonal variations were previously unknown. Knowledge of activity patterns is important for standardising census methods and census data. This study quantified the diurnal pattern in haul-out behaviour of Weddell seals at the Vestfold Hills from October 1994 to March 1995. Sequential counts of seals on the ice showed that, between 0900 and 1930 hours, seal abundance differed up to 95%. Fewer seals were hauled out in the morning than in the afternoon. The maximum numbers of seals were hauled out at the warmest time of day. The diurnal cycle was less pronounced in the breeding season than in the moulting season. The findings indicated the importance of censusing Weddell seals after 1430 hours and before 1700 hours local time, especially in the moulting season. Correction factors are given for month and time of day. Received: 27 November 1996 / Accepted: 1 May 1997     

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.     

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.     

The diel hauling-out cycle of harbour seals in an open marine environment: correlates and constraints

A colony of harbour seals in the Pacific north-west was monitored over two years, concurrent with a variety of environmental variables. Regression models described diel hauling-out activity as: i) a photoperiodic cycle; ii) a function of other environmental factors; or iii) a cycle modified by environmental constraints. Throughout the year, the number of seals on shore followed a diel pattern with a midday peak. Seals hauled-out in lower numbers in winter than in summer, and for a smaller proportion of the day (although for about the same proportion of the photoperiod). During the annual moult, numbers hauled were elevated around the clock, and the midday peak was skewed to late afternoon/early evening. Models that defined hauling-out in terms of environmental factors were significant, but did not fit the data as well as models based on photoperiod. The strongest environmental correlates (such as tidal height) owed much of their explanatory power to artefactual similarities with the photoperiodic cycle. Four general conditions are presented which, if met, should always result in a diel hauling-out cycle with a midday peak. The most fundamental of these involves a proposed 'cost of immersion'which motivates pinnipeds to haul-out when not foraging. Two likely candidates for such a cost involve risk from aquatic predators and the energetic expense of sleeping while immersed.     

Factors Affecting Haul-Out Behavior of Harbor Seals (Phoca vitulina) in Tidewater Glacier Inlets in Alaska: Can Tourism Vessels and Seals Coexist?

Large numbers of harbor seals (Phoca vitulina) use habitat in tidewater glaciers in Alaska for pupping, breeding, and molting. Glacial fjords are also popular tourist destinations; however, visitation by numerous vessels can result in disturbance of seals during critical life-history phases. We explored factors affecting haul-out behavior of harbor seals at a glacial site frequented by tourism vessels. In 2008-10, we deployed VHF transmitters on 107 seals in Endicott Arm, Alaska. We remotely monitored presence and haul-out behavior of tagged seals and documented vessel presence with time-lapse cameras. We evaluated the influence of environmental and physical factors on the probability of being hauled out, duration of haul-out bouts, and as factors associated with the start and end of a haulout. Location, season, hour, and interactions of location by year, season, hour, and sex significantly influenced haul-out probability, as did ice, weather, and vessels. Seals were more likely to be hauled out with greater ice availability during the middle of the day, and less likely to be hauled out if vessels were present. Cruise ships had the strongest negative effect; however, most vessel types negatively affected haul-out probability. Haul-out duration was longest in association with starting on incoming tides, clear skies, no precipitation, occurring in the middle of the day, and ending in the late afternoon or evening. End of haulouts was associated with increasing cloud cover, low ice availability, and vessel presence; large-sized tourism vessels or all-vessel-types combined were significant predictors of ending a haul-out bout. Probability of being hauled out was highest in June, during pupping season. Potential disturbances of harbor seals could be reduced, enabling longer resting times for seals and fewer interruptions for nursing pups, if vessels focused the majority of visits to glacial habitat to before or after the hours of 08:00-17:00 or, less optimally, 09:00-16:00.     

Mark-resight estimates of seasonal variation in harbor seal abundance and site fidelity

Monitoring trends in abundance of pinnipeds typically involves counting seals at terrestrial haul-out sites during the breeding season. Counts of seals made at other times of the year are typically lower; however, it is often unknown whether this is because fewer animals are present or whether lower counts simply reflect a reduction in haul-out probability. Here we illustrate how photo-identification data from an individual-based study of harbor seals (Phoca vitulina) can be used to estimate seasonal variation in abundance and site fidelity. Monthly data collected over a two-year period were analyzed using a mark-recapture mark-resight model accounting for individuals transitioning between observable and unobservable states. Levels of site fidelity were high throughout the year and abundance estimates showed no seasonal pattern. This suggests that individual seals used haul-out sites to the same extent throughout the year, and that peaks in counts during the breeding season are a result of seasonal changes in haul-out probability. The results of this study have implications for understanding population sub-structuring, gene flow and disease spread.     

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.     

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.     

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.     

Seasonal variation in the diet of common seals (Phoca vitulina) in the Moray Firth area of Scotland

Seasonal variation in the diet of common seals ( Phoca vitulina ) in the Moray Firth, north-east Scotland, was determined from analysis of faecal samples collected at haul-out sites during each month of 1988. Data on diet of common seals in 1987 are also presented. Limitations of the methods available for quantification of diet are discussed. Although some of the observed variation in diet from month to month may reflect changes in the sampling regime, a clear seasonal pattern was apparent, with clupeids predominating in the winter and sandeels in the summer. The trends observed are consistent with opportunistic feeding on the most abundant Prey.     

Habitat use by harbour seals (Phoca vitulina) in a seasonally ice-covered region, the western Hudson Bay

Over the last few decades, the period of ice cover in Hudson Bay has decreased, owing to climate warming, with breakup occurring approximately 3 weeks earlier than it did 30 years ago. The trend towards lengthening of the open water season has led to speculation that ringed seal numbers would decline, but then harbour seals might become numerous enough to replace ringed seals in the diet of polar bears. The movement patterns of 18 harbour seals equipped with satellite-linked transmitters in the Churchill River estuary (western Hudson Bay) were examined, as well as the dive behaviour of 11 of these seals. During the ice-free period, seals followed a general central place-foraging strategy, making repeated trips between their haul-out site in the Churchill River estuary and nearshore areas (<20 km) near the river mouth and estuary. Seal behaviour changed significantly as ice started to form along the coast of western Hudson Bay: animals remained significantly farther from the Churchill River haul-out site and from the coast and performed longer and deeper dives. However, throughout the entire tracking period, whether ice was present or not, all animals restricted their movements to a narrow band of shallow coastal waters (<50 m depth) along a 600-km stretch of the western Hudson Bay coastline, centred on the Churchill River estuary haul-out site. This natural self-limitation to nearshore shallow waters could restrict the potential for the population to increase in size and replace ringed seals as a primary energy resource for polar bears.