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.     

Modeling the effects of El Niño, density-dependence, and disturbance on harbor seal (Phoca vitulina) counts in Drakes Estero, California: 1997–2007

Harbor seal ( Phoca vitulina ) haul-out site use may be affected by natural or anthropogenic factors. Here, we use an 11-yr (1997–2007) study of a seal colony located near a mariculture operation in Drakes Estero, California, to test for natural (El Niño-Southern Oscillation (ENSO), density-dependence, long-term trends) and anthropogenic (disturbance or displacement related to oyster production activities) factors that may influence the use of haul-out subsites. Annual mariculture related seal disturbance rates increased significantly with increases in oyster harvest ( r _s= 0.55). Using generalized linear models (GLMs) ranked by best fit and Akaike's Information Criteria, ENSO and oyster production (as a proxy for disturbance/displacement) best explained the patterns of seal use at all three subsites near the mariculture operations, with effects being stronger at the two subsites closest to operations. Conversely, density-dependence and linear trend effects poorly explained the counts at these subsites. We conclude that a combination of ENSO and mariculture activities best explain the patterns of seal haul-out use during the breeding/pupping season at the seal haul-out sites closest to oyster activities.     

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.     

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.     

Spatial variation in foraging behaviour of a marine top predator (Phoca vitulina) determined by a large-scale satellite tagging program

The harbour seal (Phoca vitulina) is a widespread marine predator in Northern Hemisphere waters. British populations have been subject to rapid declines in recent years. Food supply or inter-specific competition may be implicated but basic ecological data are lacking and there are few studies of harbour seal foraging distribution and habits. In this study, satellite tagging conducted at the major seal haul outs around the British Isles showed both that seal movements were highly variable among individuals and that foraging strategy appears to be specialized within particular regions. We investigated whether these apparent differences could be explained by individual level factors: by modelling measures of trip duration and distance travelled as a function of size, sex and body condition. However, these were not found to be good predictors of foraging trip duration or distance, which instead was best predicted by tagging region, time of year and inter-trip duration. Therefore, we propose that local habitat conditions and the constraints they impose are the major determinants of foraging movements. Specifically the distance to profitable feeding grounds from suitable haul-out locations may dictate foraging strategy and behaviour. Accounting for proximity to productive foraging resources is likely to be an important component of understanding population processes. Despite more extensive offshore movements than expected, there was also marked fidelity to the local haul-out region with limited connectivity between study regions. These empirical observations of regional exchange at short time scales demonstrates the value of large scale electronic tagging programs for robust characterization of at-sea foraging behaviour at a wide spatial scale.     

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 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.     

Satellite tracking of ringed seals Phoca hispida off northwest Greenland

We monitored movements and haul-out patterns of four ringed seals Phoca hispida , off Northwest Greenland between 5 June and 31 October 1988 using the Argos Data Collection and Location System When the seals were hauled out on fast ice their locations were accurately determined, but when they were at sea, few accurate locations were obtained, evidently because these seals spent little time at the surface between dives The seals remained within the fjord where they were tagged, and hauled out often to early July Thereafter, as fast-ice disappeared, they dispersed widely and spent less time hauled out Time of day had no significant effect on haul-out patterns Haul-out periods declined significantly from June to August and increased in September-October Satellite contact with one seal was lost after 16 d while the seal was still in the fjord in late June One seal travelled over 200 km southwest and was located 4 July in offshore waters of Smith Sound 30 d after instrumentation Another seal moved southeast along the Greenland coast where contact was lost after 49 d on 23 July The fourth seal moved north along the Greenland coast, hauled out regularly on ice, and returned south along the coast in late September and October after 181 d of contact with the satellite     

Sex differences in the nursing relationship between mothers and pups in the Atlantic harbour seal, Phoca vitulina concolor

Sex differences in the dynamics of the relationship between harbour seal mothers and their pups were investigated. An index of pair proximity governance suggested that pups became increasingly responsible for maintaining the distance between the pair over the course of the nursing period. Male pups demonstrated a stronger tendency to control initiation distance than female pups. A Nursing Index utilizing suckling bout initiations and terminations revealed that, as weaning approached, pups became progressively more responsible for the continuance of nursing. Male pups exhibited more control than female pups, primarily through initiating a greater portion of the bouts. On-teat times averaged 4.08 min per bout, with no significant differences between male and female pups. Male pups, however. nursed a greater portion of haul-out time due to higher bout frequencies obtained through more bout initiations. Mothers did not preferentially reject attempts to nurse by pups of either sex. These results suggest that the tenet of parental investment theory suggesting male pups enjoy preferential investment may be misleading. In harbour seals. male pups seem actively to seek out resources, as opposed to any inherent strategy on the par1 of the mother.     

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.     

A Bayesian estimate of harbour seal survival using sparse photo-identification data

Survival rates have rarely been estimated for pinniped populations due to the constraints of obtaining unbiased sample data. In this paper, we present an approach for estimating survival probabilities from individual recognition data in the form of photographic documentation of pelage patterns. This method was applied to estimate adult (age 2+) survival for harbour seals in the Moray Firth, NE Scotland. An astronomical telescope was used to obtain digital images of individual seals, and high-quality images were used to document the annual presence or absence of individuals at a single haul-out site over a 4-year period. A total of 95 females, 10 males and 57 individuals of unknown sex were photographically documented during the study period. Survival and recapture probabilities were estimated using Jolly–Seber mark–recapture models in a Bayesian statistical framework. Computer-intensive Markov Chain Monte Carlo methods were used to estimate the probability distributions for the survival and recapture probabilities, conveying the full extent of the uncertainty resulting from unavoidably sparse observational data. The deviance information criterion was used to identify a best-fitting model that accounted for variation in the probability of capture between sexes, with constant survival. The model estimated adult survival as 0.98 (95% probability interval of 0.94–1.00) using our photo-identification data alone, and 0.97 (0.92–0.99) with the use of an informative prior distribution based on previously published estimates of harbour seal survival. This paper represents the first survival estimate for harbour seals in the UK, and the first survival estimate using photo-identification data in any species of pinniped.     

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     

Breeding habitat and lair structure of the ringed seal (Phoca hispida ladogensis) in northern Lake Ladoga in Russia

The breeding habitat and lair structures of the ringed seal (Phoca hispida ladogensis) were studied by snowmobile expeditions in northern Lake Ladoga, Russia, during 1996 to 1999. Mostly flat ice and no large pack ice existed in northern Lake Ladoga during this study. All birth lairs and 88% of haul-out lairs were located in snowdrifts in the shorelines of islands or islets and 12% of the haul-out lairs occurred in pressured ice ridges. Some large haul-out lairs had been used communally by the seals. A total of 15% of the lairs observed had been attacked or marked by red fox (Vulpes vulpes), wolf (Canis lupus) or unidentified canines. The colour of the lanugo fur of the Ladoga ringed seal pup appeared to be dark grey. We conclude that the northern part of Lake Ladoga is an active breeding area for the ringed seal, which attests conservation needs for this area. Accepted: 18 September 2000     

DIVING BEHAVIOR OF THE SAIMAA RINGED SEAL (PHOCA HISPIDA SAIMENSIS NORDQ.)

The activity and diving patterns of four adult Saimaa ringed seals ( Phoca hispida saimensis , a landlocked subspecies living in Lake Saimaa, Finland) were examined during spring, summer, and autumn by the use of VHF-transmitters. Over 17,000 dives were registered. The duration of the dives and diving patterns differed among individuals. The mean duration of dives increased from spring to autumn; e.g. , in one individual the mean dive duration increased from 6 min in June to 10.5 min in October. The haul-out periods of one individual in May to early June made up 46.2% of its total activity budget, but in another individual in July to August the haul-out periods made up only 11% of the budget and the seal was submerged for 80% of the time. Periods of successive long duration dives (>10 min) were observed in three individuals in summer and autumn. The longest dive measured was 23 min. The duration of the periods containing long dives was often over three hours (maximum six hours) and the mean duration of the dives about 15 min. These long duration dives are assumed to be aerobic resting dives. Generally, the dives of the Saimaa ringed seal appear to be of longer duration than previously assumed.     

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.     

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.     

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.     

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.     

Apparent decline in the harbour seal Phoca vitulina population near Hvaler, Norway, following an epizootic

From 1986-90 we tagged 98 harbour seal pups at Hvaler, Norway Fifteen were recovered later the same year of which 10 were recovered in 1988, when the seal population was infected by a morbillivirus The epizootic in 1988 resulted in a population decline of c 75% Predicted birth weight did not vary significantly before and after the epizootic Newborn harbour seals have antibodies against the morbillivirus two years after the outbreak of the disease     

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.