LONG-TERM TRENDS IN HARBOR SEAL NUMBERS AT TUGIDAK ISLAND AND NANVAK BAY, ALASKA

We conducted land‐based counts of harbor seals (Phoca vitulina richardii) and collected related environmental data at Tugidak Island (Gulf of Alaska, 1994–2000) and Nanvak Bay (Bristol Bay, 1990–2000) to estimate population trends and identify factors influencing counts. At Tugidak Island, the seal population declined substantially during molting from 1976 through the 1980s, stabilized in the early 1990s, and increased at a moderate rate (3.4%/yr, CI: 1.0%–5.8%) from 1994 to 2000. Pups and all seals ashore during pupping increased at higher annual rates of 5.4% (CI: 2.2%–8.8%) and 8.3% (CI: 4.5%–12.3%) from 1994 to 2000 at Tugidak Island. At Nanvak Bay seals declined in abundance between 1975 and 1990 but increased during the 1990s at 9.2%/yr (CI: 7.2%–11.3%) during pupping and 2.1%/yr (CI: 0.6%–3.6%) during molting. Date and time‐of‐day were significant covariates in all analyses. Factors that led to declines at Tugidak Island and Nanvak Bay have seemingly abated sufficiently such that these populations are currently increasing, though still greatly reduced from the 1970s. Index sites are useful adjuncts to aerial surveys, providing survey‐related information not always available from aerial counts, which is useful in survey design and data analysis.     

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.     

Earlier pupping in harbour seals,Phoca vitulina

The annual reproductive cycle of most seal species is characterized by a tight synchrony of births. Typically, timing of birth shows little inter-annual variation. Here, however we show that harbour seals Phoca vitulina from the Wadden Sea (southeast North Sea) have shortened their yearly cycle, moving parturition to earlier dates since the early 1970s. Between 1974 and 2009, the birth date of harbour seals shifted on average by −0.71 d yr⁻¹, three and a half weeks (25 days) earlier, in the Dutch part of the Wadden Sea. Pup counts available for other parts of the Wadden Sea were analysed, showing a similar shift. To elucidate potential mechanism(s) for this shift in pupping phenology, possible changes in population demography, changes in maternal life-history traits and variations in environmental conditions were examined. It was deduced that the most likely mechanism was a shortening of embryonic diapause. We hypothesize that this could have been facilitated by an improved forage base, e.g. increase of small fishes, attributable to overfishing of large predator fishes and size-selective fisheries.     

Large increases of harp seals (Phoca groenlandica) and hooded seals (Cystophora cristata) on Sable Island, Nova Scotia, since 1995

Beach surveys for harp (Phoca groenlandica) and hooded (Cystophora cristata) seals documented a dramatic increase in their numbers on Sable Island in the mid-1990s. From late 1994 to 1998, 1,191 harp and 870 hooded seals, mostly young animals, were recorded on the island whereas, in the 1980s, no more than 5 animals of both species were observed each year. Of the 2,061 harp and hooded seals examined, 41.7% were found alive, 26.7% were killed by sharks, and 31.6% were found dead but intact. This increase in numbers of harp and hooded seals on Sable Island, which is south of their historic northern range, is consistent with the recent increase of extralimital occurrences of these species along the east coast of North America. However, the large number of seals recorded in this study provides more information on their demography than has previously been possible.     

Effects of maternal age and condition on parturition and the perinatal period of Antarctic fur seals

The effect of maternal age and condition on the date of parturition and the duration of the perinatal period of Antarctic fur seals at Bird Island, South Georgia, were investigated over three consecutive breeding seasons. Females rear young during a four-month lactation period in a highly seasonal but predictable environment. Although females may first pup at three years of age, they did not attain full adult size until six years of age; older females (≥ 6 years) tended to be heavier, longer, and in better condition than younger females (3–5 years). Older females returned to breeding beaches earlier and could occupy the most suitable pupping sites, and gave birth when densities of animals on the beaches were low (i.e. more favourable for pup survival). Females that arrived earlier were able to remain ashore longer with their pups prior to departing on their first foraging trips but this was unrelated to either maternal age or condition. Younger females returned later in the pupping season, possibly as a result of late implantation due to smaller energy reserves than older and larger females. In 1990 all females arrived late, were in poorer condition, gave birth to lighter pups, and had shorter perinatal periods. This suggests that not only was implantation late but that females returned to an area of low food availability prior to parturition.     

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.     

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.     

Variation in breeding phenology provides insights into drivers of long-term population change in harbour seals

Phenological trends provide important indicators of environmental change and population dynamics. However, the use of untested population-level measures can lead to incorrect conclusions about phenological trends, particularly when changes in population structure or density are ignored. We used individual-based estimates of birth date and lactation duration of harbour seals (Phoca vitulina) to investigate energetic consequences of changes in pupping phenology. Using generalized linear mixed models, we first demonstrate annual variation in pupping phenology. Second, we show a negative relationship between lactation duration and the timing of pupping, indicating that females who pup early nurse their pups longer, thereby highlighting lactation duration as a useful proxy of female condition and resource availability. Third, individual-based data were used to derive a population-level proxy that demonstrated an advance in pupping date over the last 25 years, co-incident with a reduction in population abundance that resulted from fisheries-related shootings. These findings demonstrate that phenological studies examining the impacts of climate change on mammal populations must carefully control for changes in population density and highlight how joint investigations of phenological and demographic change provide insights into the drivers of population declines.     

TRENDS IN ABUNDANCE OF ALASKA HARBOR SEALS, 1983–2001

We estimated trends in abundance of harbor seals ( Phoca vitulina richardsii ) using over dispersed, multinomial models and counts obtained during aerial surveys conducted during 1983–2001 in the Ketchikan, Sitka, Kodiak, and Bristol Bay areas of Alaska. Harbor seal numbers increased significantly at 7.4%/yr during 1983–1998 and 5.6%/yr during 1994–1998 in the Ketchikan area, and 6.6%/yr during 1993–2001 in the Kodiak area. Counts were stable (trends not significant) during 1984–2001 (0.7%/yr) and 1995–2001 (-0.4%/yr) in Sitka, and during 1998–2001 (-1.3%/yr) in Bristol Bay. The influence of covariates ( e.g. , survey date, tide height) on trend estimates was significant and varied among areas and across years, demonstrating the need to include covariates in statistical analyses to accurately estimate trend. Our increasing trend estimate for Kodiak represents the first documented increase in harbor seal numbers over a relatively expansive area in the Gulf of Alaska. However, the trend for the Gulf of Alaska stock is equivocal due to the continued decline in Prince William Sound. Similarly, the trend for the Southeast Alaska stock is equivocal based on our increasing (Ketchikan) and stable (Sitka) trend estimates, and a recent decline reported for Glacier Bay. The Bering Sea stock appears stable after a period of possible decline.     

Haulout behaviour of High Arctic harbour seals (<Emphasis Type="Italic">Phoca vitulina vitulina</Emphasis>) in Svalbard,Norway

Haulout behaviour of harbour seals, living at the northern limit of their distributional range on Svalbard, Norway, was investigated from June to August 2000 using a combination of low-tide counts performed during boat surveys, hourly counts through 12- or 24-h cycles at specific haulout sites, and telemetric data from 37 VHF-tagged seals. The largest aggregations of seals were found at Skarvnes, a site where numbers increased steadily through the summer, reaching a peak during the moulting period in August. At this site, season/date, time of day, tidal state and temperature all significantly influenced the number of animals ashore. At the second most frequented haulout site, at Sørøya, season/date, time of day, temperature and cloud cover significantly affected the number of seals using the site. Pups were found predominantly at Sørøya (7.8 pups±6.3 SD, N=53 counts); they were less common at Skarvnes (1.0 pups±0.2 SD, N=95 counts). Haulout patterns varied by age and sex class in accordance with the demands of lactation, mating and moult. Our limited data on mother-pup pairs suggest that they are closely associated during the nursing period, spending approximately 50% of their time hauled out together. Post lactation, most adult females left haulout areas for periods of up to several days. The haulout behaviour of adult males suggested that they adjusted their behaviour to follow female distribution and movement patterns during the breeding period. Most juveniles and adults of both sexes stayed ashore for prolonged periods during moulting, which took place first in juveniles, then in adult females and finally in adult males. The results of our study show that the basic haulout behaviour patterns of harbour seals at Svalbard are similar to this species behaviour at lower latitudes.     

Dispersion and site fidelity of breeding female grey seals (Halichoerus grypus) on North Rona, Scotland

Female grey seals ( Halichoerus grypus ) formed breeding aggregations on the island of North Rona, Scotland. Aggregations of females were associated particularly with gullies leading from the sea, leaving large areas of available space unoccupied. Changes in the degree of aggregation of females during the breeding season were similar in 1987, 1988 and 1989. Pronounced aggregations occurred in the early and late parts of each breeding season.
Of 67 breeding females marked in 1985, 62 (93%)) returned to N. Rona to breed in at least one season up to 1989, but 18 (27%) were present in all five years. Females came ashore up to 14 days before giving birth and 82% were observed first in the vicinity of their subsequent pupping site. Between 1985 and 1989, marked females which returned were faithful to their previous pupping sites, even when the previous pup had died. There was no evidence of a gradual change in the location of individual pupping sites over time. This pupping site fidelity may generate aggregations whose location, timing and composition is predictable.     

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.     

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.     

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

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

GRAY WHALE CALF PRODUCTION 1994–2000: ARE OBSERVED FLUCTUATIONS RELATED TO CHANGES IN SEASONAL ICE COVER?

We conducted shore-based sighting surveys to estimate the number of northbound migrating gray whale calves passing Piedras Blancas, California, for seven consecutive years (1994–2000). In addition, we conducted aerial surveys to determine offshore distribution of the migration in 1994 and 1995, measured day/night migration rates with thermal sensors in 1994–1996, and maintained concurrent replicate watches near the peak of each migration to estimate the proportion of the cow/calf pairs missed by the standard watch team. During good weather, we counted 325, 194, 407, 501, 440, 141, and 96 calves during 1994–2000, respectively. Correcting these counts for periods not on watch and for calves missed, produced final estimates of 945 calves (SE = 68.21) for 1994, 619 calves (SE = 67.19) for 1995, 1,146 calves (SE = 70.67) for 1996, 1,431 calves (SE = 82.02) for 1997, 1,388 calves (SE = 91.84) for 1998, 427 calves (SE = 41.10) for 1999, and 279 calves (SE = 34.79) for 2000. Calf production indices (calf estimate/total population estimate) are 4.2%, 2.7%, 4.8%, 5.8%, 5.5%, 1.7%, and 1.1% for the years 1994–2000, respectively. Fluctuations in calf production over this time period were positively correlated with the length of time that primary feeding habitat was free of seasonal ice during the previous year.     

Effects of the lunar cycle on the Galápagos fur seal,Arctocephalus galapagoensis

Summary During the 1977 and 1979 reproductive periods of the Galápagos fur seals a census taken in the mornings and evenings at Cabo Hammond, Fernandina, showed a marked, synodic lunar rhythm in numbers of animals ashore. About twice as many fur seals were ashore at full moon than at new moon. By use of two independent Fourier analysis methods, the curve of the morning counts is shown to lag 15°–20° of the lunar month behind the curve of the evening counts. The lunar effect is demonstrated for males, females, and immatures. The rhythm is also seen is demonstrated for males, females, and immatures. The rhythm is also seen in attendance data from 13 individually marked females, all but one nursing young. Reproductive events show the lunar rhythm much less markedly than do numbers ashore. This and the clear rhythm in immature numbers make it very likely that the rhythm is a year-round phenomenon, independent of reproduction.There is no reason to assume that fur seals stay on land during moonlit nights especially for social interaction. It is then hypothesized that fur seals avoid moonlight at sea. If so, the peak of numbers ashore at full moon and the negative phase angle difference of the evening curve against the morning curve can be explained with the shift, and the varying duration and brightness, of the moonlit part of the night over the lunar cycle. Two hypotheses which might account for this moonlight avoidance are discussed: (1) predator (shark) avoidance and (2) varying feeding efficiency of the fur seals due to the influence of moonlight on the vertical distribution of prey.     

Grey seal, Halichoerus grypus, energetics: females invest more in male offspring

Grey seal females transfer large amounts of energy to their pups during the brief lactation period. The costs of lactation have been measured using weight changes of mother and pup pairs. Large females come ashore to give birth earlier in the season and lose weight more rapidly than smaller females. The sex ratio of Grey seal pups born is skewed towards males in the early part of the breeding season. Male pups are larger at birth and gain weight more rapidly than female pups, and their mothers show a correspondingly faster rate of weight loss than mothers of female pups. The energy costs of gestation and lactation to a Grey seal mother are 31 GJ for male pups and2–8 GJ for female pups. Males are therefore 10% more costly in energy terms to raise to weaning. Because, on average, large females arrive at breeding sites before smaller animals, biased results on weight changes would be obtained from methods which do not use repeated weighings. We suggest that the high efficiencies of lactation estimated for Harp seals compared with other phocid seals could be accounted for by such a bias.     

Winter distribution of harp seals (Phoca groenlandica) off eastern Newfoundland and southern Labrador

The winter distribution of Newfoundland harp seals (Phoca groenlandica) was determined using sighting data collected during January and February from 1991 to 1995 aboard research vessels that covered the northeastern continental shelf between 46–55°N and 47–54°W. Data were standardized for effort and sighting conditions. Visual appraisals of data were made using a Geographical Information System. In contrast to historical perceptions, offshore areas such as the northern part of the Grand Banks (48–49°N and 49–51°W) appear to be extremely important to wintering harp seals. Southeastern shifts in distribution appear to have occurred since the early 1990s, particularly between the 1991–1993 and 1994–1995 periods. This southern shift in range agrees with the recent increase in extralimital occurrences of harp seals along the North American east coast. Concurrent changes in environmental conditions suggest that physical and biological factors may influence the distribution of this population. Accepted: 22 May 2000     

Climatic regime shifts and decadal-scale variability in calanoid copepod populations off southern California

Decadal‐scale climatic regimes and the shifts between them have important impacts on marine ecosystems. Climatic regime shifts have been observed or hypothesized in the North Pacific basin in 1976–77 and 1989. This paper examines long‐term (1951–99) trends in calanoid copepod populations off southern California, and the evidence for responses to regime shifts. Most of the species of calanoid copepod that were analysed underwent one or more step changes during the 49 years covered by the study. All but one of these changes occurred in five periods: the late 1950s, late 1960s, mid‐1970s, early 1980s and around 1990. The late 1960s changes are considered to be artifacts of an increase in sampling depth. Strong El Niño conditions affected California waters during the late 1950s and early 1980s. The step changes of the mid‐1970s and late 1980s to early 1990s may have been responses to regime shifts or other climatic events. 28% of the species and subspecies responded to the 1976–77 event, all increasing in abundance. Another 28% of the copepod categories underwent step changes around 1990, most decreasing. Evidence for regime shifts in the hydrographic variables that were examined is mixed. The 10‐m temperature increased in the mid‐1970s. Abrupt changes in variables around 1990 were short‐lived. However, the population responses around 1990 and to the El Niños of the late 1950s and early 1980s indicate that some species of calanoid copepods may respond on longer time scales to environmental conditions that persist only a few years.     

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.