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.     

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

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

INCREASED VIGILANCE TOWARD UNFAMILIAR HUMANS BY HARBOR (PHOCA VITULINA) AND GRAY (HALICHOERUS GRYPUS) SEALS

Two pairs of harbor ( Phoca vitulina ) and three pairs of gray ( Halicboeruls grypus ) seals were exposed to one of three human handlers for 15 min, twice a day, for a total of six sessions. Following habituation to the familiar handler, animals were then exposed to a novel human for 7 min, and then retested for 7 min with the familiar human. In all cases, animals responded to the unfamiliar human with increased vigilant behavior, i. e., they spent more time oriented towards the unfamiliar handler during the first 2 min of the test session than during the same interval of either the final habituation session or the retest with the familiar human ( P = 0.03 in all cases). There was also a tendency for seals to contact the familiar handler with their noses more rapidly than the novel human ( P = 0.06). These results support the hypothesis that phocid seals are capable of discriminating between individual humans in their environment, setting the stage for human-based Pavlovian conditioning.     

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.     

GROWTH AND DEVELOPMENT IN FREE-RANGING HARBOR SEAL (PHOCA VITULINA) PUPS FROM SOUTHERN BRITISH COLUMBIA, CANADA

Harbor seals ( Phoca vitulina ) are small pinnipeds that are widely distributed throughout the temperate coastal regions of the Atlantic and Pacific oceans. We determined birth mass, neonatal growth rates, weaning age, and weaning mass of NE Pacific harbor seals ( P. v. richardsi ) during a capture-recapture study that spanned the nursing period (Sidney Island, British Columbia, Canada). Of 46 harbor seal pups initially captured, 28 were classified as newborns ( i. e. , < 24 h old). Mean body mass of newborns was 11.2 ± SE 0.31 kg. Pups were individually tagged and recaptured throughout the nursing period. Average daily mass gain during the nursing period was 394 ± 26 g. Mean birth mass of males did not differ significantly from females, although pups found with fetal pelage (lanugo) (21.4% of all newborns) were smaller at birth (9.8 ± 0.44 kg) than non-lanugo pups (11.6 ± 0.33 kg). Mean weaning mass was estimated at 23.6 ± 1.2 kg at a mean weaning age of 32 d ± 1.5 d. While birth and weaning masses differed little from the published data for offshore Sable Island harbor seals ( P. v. concolor ), British Columbia harbor seals are characterized by half the daily mass gain and a longer nursing period.     

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.     

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.     

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.     

THE DISTRIBUTION, ABUNDANCE AND SELECTED PREY OF THE HARBOR SEAL, PHOCA VITULINA CONCOLOR, IN SOUTHERN NEW ENGLAND

The seasonal distribution and abundance of harbor seals occurring south of Maine were documented by counting the number of seals at traditional haulout locations. The average number of seals counted during each survey in Massachusetts and New Hampshire was 3,560 ± 255 (95% CI), 1983–1987. The maximum number of seals counted on any individual survey was 4,736 individuals. Fifty percent of all the surveys since January 1985 have resulted in counts greater than 4,000 seals reflecting a 27% increase in the abundance of seals in our study area since that date. Seventy-five percent of the seals in southern New England are located at haulout sites on Cape Cod and Nantucket Island. The largest aggregation of seals in the eastern United States occurs mid-winter at Monomoy Island and adjacent shoals. A single high count of 1,672 seals occurred at this site during the study period. An additional 271–374 seals were also counted in Rhode Island, Connecticut and eastern Long Island Sound during surveys conducted in March 1986 and 1987. The American sandlance Ammodytes americanus was the single dominant prey item of harbor seals in waters adjacent to Cape Cod based on the modified frequency of occurrence of each prey species in scat samples collected from three haulout sites on Cape Cod between 1984–1987. During January and February sandlance was the near exclusive prey item at Monomoy (99%, n= 80). During March and April, the frequency of Atlantic herring Clupea harengus increased in the scat samples at this site. Regional differences in the diet of seals reflect distinct prey communities throughout the study area. Since 1986, the percent occurrence and importance of sandlance in the diet of seals has decreased, reflecting an overall decrease in abundance of this prey species in waters adjacent to Cape Cod. In spite of fluctuations in abundance, and regional differences in the diet of seals throughout the study area, sandlance still comprised a minimum 55% of the total prey species of harbor seals throughout the study area.     

FEEDING METHOD AFFECTS OTOLITH DIGESTION IN CAPTIVE GRAY SEALS: IMPLICATIONS FOR DIET COMPOSITION ESTIMATION

Analysis of prey structures from scats has become widely used for assessing the diet of free-ranging seals. Captive feeding experiments have played an important role in the development of scat analysis but methodology has varied. We investigated whether using an experimental otolith-carrier species or in situ otolith experimental prey to feed haddock, plaice, and sand eel otoliths to two captive gray seals affected the amount of otolith digestion, and therefore our ability to estimate fish size and diet composition. Otolith recovery rates varied among the three prey species, but were not affected by feeding method. Otoliths were more digested in carrier experiments; resulting digestion coefficients were greater than those from in situ experiments. As a result, while the lengths of whole fish fed experimentally could be accurately predicted by applying in situ digestion coefficients, the lengths of these fish could not be accurately predicted using carrier digestion coefficients. Consequently, the estimated proportion by weight of each species in the diet can change markedly when carrier, as opposed to in situ , digestion coefficients are used. The implications for diet composition estimation are significant.     

REDUCED POPULATION GROWTH OF GRAY SEALS AT SABLE ISLAND: EVIDENCE FROM PUP PRODUCTION AND AGE OF PRIMIPARITY

Pup production on Sable Island, Nova Scotia, has been increasing exponentially since the early 1960s and by 1997 Sable Island was the largest gray seal colony worldwide. Using an aerial photographic survey, as in previous years, we estimated pup production in January 2004 to determine if this exponential rate of increase had continued. A total of 33,268 pups was counted on the color positives. When corrected for the proportion pups missed on the imagery (1.106 for the 12th; 1.527 on the 13th), the proportion of pups that died prior to the survey (0.031), and the proportion of pups born before the survey (east colony 0.966, west colony 0.962), estimated total pup production was 41,500 with SE = 4,381. The 2004 estimate indicates that pup production on Sable Island has continued to increase, but suggests that the rate of increase ( r ) may have declined (0.070 compared to previous 0.128). Females from the 1998–2000 cohorts were about 16 times less likely to give birth for the first time at age 4 yr and more than twice as likely at age 6 yr compared to those in the mid-late 1980s. The new estimate of pup production and observed changes in age of primiparity provide the first indication of changes in vital rates of this population. However, additional estimates of pup production and vital rates are needed to confirm this conclusion and to investigate the underlying mechanisms.     

THE MAXIMUM RANGE OF VOCAL COMMUNICATION IN AIR BETWEEN A HARBOR SEAL (PHOCA VITULINA) PUP AND ITS MOTHER

Abstract: The transmission losses with distance of four pure tones (0.5, 1, 2, 4 kHz) were measured in the air 9 cm above the surface of the coastal waters in the Bay of Fundy. The study was conducted between May and July 1992, on warm days with low winds. The measured transmission losses were as much as 11 dB less than predicted by spherical spreading (6 dB/distance doubled), at 400 m (0.5 kHz). This enhanced sound transmission is probably due to the air temperature profile which increases with height above the water's surface. Such a profile causes sound waves to refract towards the water, thereby reducing spreading losses. High-frequency sound absorption negates enhanced transmission at 4 kHz, at distances over 500 m. On days with low winds and low ambient noise levels, a seal pup calling at 90 dB re 20, μPa at 0.5 kHz should be detectable by the mother up to 1 km away, and may be audibly recognizable up to 140 m away.     

ESTIMATION OF AGE-SPECIFIC PROBABILITIES OF FIRST RETURN AND ANNUAL SURVIVAL RATES FOR THE MALE GRAY SEAL (HALICHOERUS GRYPUS) ON SABLE ISLAND FROM CAPTURE-RECAPTURE DATA

We used a longitudinal capture-recapture study to estimate the age-specific probabilities of first return to the breeding colony and annual survival rates for male gray seals ( Halichoerus grypus ), based on resightings of seals branded as young on Sable Island. We estimated that the average age of first returns for seals born in 1969–1970 to be 9.1 (SE 0.4) yr; for seals born in 1973–1974 it is estimated to be 9.8 (SE 0.2) yr. The estimated annual survival rate of these males was estimated to be 0.976 (SE 0.003).     

HARBOR SEAL (PHOCA VITULINA) PREDATION ON SEINED SALMONIDS IN THE LOWER KLAMATH RIVER, CALIFORNIA

Feeding activity of harbor seals ( Phoca vitulina ) was monitored while the California Department of Fish and Game seined and tagged migrating adult salmonids between 1984 and 1988. The number of predations observed each week of observation was significantly correlated with the number of fish seined during that week. There was a significantly higher number of predations observed on days when seining took place than on days when no seining occurred. Our observations suggest that most, if not all, predations we observed on days when seining occurred involved fish that had been recently seined and released. The estimated percentage of seined fish taken by seals was relatively constant over the five years of the study, ranging from 3.1% to 5.5%. Various strategies for reducing the level of predation on seined salmonids in the lower Klamath River are discussed.     

BEHAVIORAL EVIDENCE OF THERMAL STRESS FROM OVERHEATING IN UK BREEDING GRAY SEALS

Gray seals ( Halichoerus grypus ) in the UK exhibit clear preferences for pupping close to pools of water on inland breeding sites. The reasons for this are unclear as seals are thought to derive all their water requirements from the metabolism of fat. Likewise, the prospect of seals overheating during the UK's cool, damp, autumnal breeding seasons, has seemed unlikely, but has not been previously explored. Here, we provide preliminary behavioral evidence of thermal stress in female gray seals breeding on the island of North Rona, Scotland. Video footage provided measures of proximity to, and the proportion of animals bathing in, pools in relation to meteorological data (temperature, mean sea-level pressure, rainfall, wind speed and direction) on four dates spread through a single breeding season. The proportion of females close to pools increased with pressure (warmer, drier conditions) and decreased on wetter days. In addition, analyses of colony-wide patterns of seal dispersion showed that females tended to be closer to pools of water at higher pressures and temperatures and at lower wind-speeds. These results provide the first evidence of thermal stress and behavioral thermoregulation through access to pools of water in a phocid breeding in temperate autumnal conditions.     

GENETIC SUBSTRUCTURE OF THE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDSI) OFF WASHINGTON, OREGON, AND CALIFORNIA

Genetic substructure among groups of Pacific harbor seals, Phoca vitulina richardsi , along the western coast of the United States was investigated using mitochondrial DNA sequences. Blood and tissue samples were removed from 86 seals inhabiting Puget Sound and the Pacific coasts of Washington, Oregon, and California. A 320 base-pair segment of the control region was amplified using the polymerase chain reaction and directly sequenced. These data indicated a high level of diversity. Thirty variable sites were found that define 47 mitochondrial haplotypes. Among groups of P. v. richardsi sampled, 5 haplotypes were shared, but most (42) were unique to a locality. Haplotypic frequency and an Analysis of Molecular Variance (A mova ) revealed significant differences ( P = 0.001) among regions. Phylogenetic analysis indicated Puget Sound seals possess unique divergent lineages not found in seals from the coasts of Washington, Oregon, and California. These lineages may represent haplotypes from north of Washington, which is consistent with late reproductive timing of harbor seals from Puget Sound.     

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.     

ASSESSMENT OF ARGOS LOCATION ACCURACY FROM SATELLITE TAGS DEPLOYED ON CAPTIVE GRAY SEALS

The Argos satellite system is commonly used to track and relay behavioral data from marine mammals, but their underwater habit results in a high proportion of locations of non-guaranteed accuracy (location classes (LC) O, A, and B). The accuracy of these locations is poorly documented in marine mammals. We assessed the accuracy of all LCs on four juvenile gray seals fitted with Argos satellite relay data loggers and held in captivity in an outdoor tank for a total of 61 seal-days. Four hundred and twenty-six locations were obtained from seals in captivity, and their latitude and longitude error was assessed before and after filtering, following MConnell et al. (1992). There was significantly more error in longitude than latitude in all LCs except I. C A. The ratio of the standard deviations of longitude : latitude ranged from 1.77 (LC 3) to 2.58 (LC 1). Filtering had very little effect on errors in LCs 3-1, but in the remaining LCs filtering resulted in error reductions ranging from 8% to 63%. In LCs O, A, and B, error reduction was greater in the 95th percentile errors, especially in longitude. The averages of the latitude and longitude 68th percentile errors and those predicted by Argos (in brackets) were 226 (150), 372 (350), and 757 (1000) m for LCs 3, 2, and 1 respectively. Both latitude and longitude errors of LCs > O were normally distributed. Both filtered and unfiltered LC A locations were of a similar accuracy to LC 1 locations, and considerably better than LC O locations.     

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.     

DIETARY AND SEASONAL INFLUENCES ON BLOOD CHEMISTRY AND HEMATOLOGY IN CAPTIVE HARBOR SEALS

It has become increasingly important to understand the relationship between dietary intake and blood chemistry and hematological profiles in pinniped populations, especially where nutritional stress has been implicated in population declines. Blood chemistry and hematology reference ranges have been commonly reported for both free‐ranging and captive harbor seals, but seasonal and dietary fluctuations are limited to only a few studies. We quantified changes in plasma metabolites and hematology values in captive harbor seals on different diets over two years. Of 29 blood chemistry and hematology variables examined in captive harbor seals, ALT, AST, GGT, creatinine, and BUN:creatinine were influenced by diet. BUN, Na⁺, Cl^?, Hct, and Hb were influenced by season. Diet did not affect hematology variables. While these data were collected on captive animals, we assume seasonal and dietary influences would be comparable in free‐ranging seals. According to this study, controlling for season during sampling becomes important when assessing blood chemistry values as an index of foraging status or health.