SEASONAL MOVEMENTS AND DIVE PATTERNS OF JUVENILE BAIKAL SEALS, PHOCA SIBIRICA

The Baikal seal ( Phoca sibirica ) is confined to Lake Baikal in southern Siberia. The breeding distribution of seals in winter, when the lake is frozen over, is fairly well known, whereas their movements and foraging behaviors have been relatively unstudied. With satellite-linked radio transmitters, we documented the movements and dive patterns of four juvenile Baikal seals from autumn through spring. The seals moved extensively in the lake, each covering minimal distances of 400–1,600 km between September and early May. They spent little time hauled out from September through May and, apparently, dived continuously. Dives were mostly to depths of lo-50 m, though a few exceeded 300 m. Most lasted between 2 and 6 mm, within theoretical aerobic dive limits, although a few exceeded 40 min. The exceptionally long dives occurred while the seals were in areas of extensive ice cover, suggesting that they were, perhaps, under ice-pilotage in search of breathing holes rather than foraging dives. Otherwise, the dive performances of these Baikal seals were, relative to body mass, similar to those of other well-studied phocids. Movements and dive patterns of seals appeared to be primarily associated with seasonal and die1 movements of their primary prey, golomyanka and sculpins, and secondarily correlated with patterns of ice formation and thaw.     

PUPPING IN THE MOST NORTHERLY HARBOR SEAL (PHOCA VITULINA)

The Norwegian high arctic archipelago Svalbard is the home of the world's northernmost population of harbor seals. Due to their limited distribution in isolated areas of the archipelago, little is known about their biology. Until now no information has been published on their pupping or whether it differs from that of harbor seals in other regions. This paper shows that the peak pupping period for harbor seals in Svalbard coincides with that in Europe, i.e., the second half of June. The pups are born with an adult-like pelt and not with a white coat.     

DIVING BEHAVIOR OF THE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDII) IN MONTEREY BAY, CALIFORNIA

Physical environment and physiological characteristics of marine mammals potentially affect the duration and depth of diving. Härkönen (1987b) proposed a hypothesis that the harbor seal would gain maximum energy by foraging at intermediate depths. To investigate this hypothesis, we studied diving behavior of the Pacific harbor seal (Phoca vitulina ricbardii) during 1995 through 1997 in Monterey Bay, California. Dive depths (n = 13,063 dives) were recorded via time‐depth recorders. Approximately 80% of recorded dives were classified as square dives (type I), which typically were associated with foraging in pinnipeds. Approximately 11% of dives were V dives (type II; 1,402 dives), and the remainder (1,225 dives) were skewed dives (type III and IV). The deepest recorded dive was 481 m, while the greatest duration was 35.25 min. Body mass explained the variability of durations of long dives for females (95th percentile; D₉₅♂=‐5.47 + 0.18 × (mass♀), r²= 0.91, 95% CI for slope = [0.08, 0.28], n= 5) and for males (D₉₅♂=‐5.86 + 0.18 × (mass♀), r²= 0.83, 95% CI for slope = [0.12, 0.24], n= 11). The large proportion of variability in deep dives, however, was explained by body mass only for males (95th percentile; Z₉₅♂=‐363.9 + 6.05 × (mass♀), r²= 0.83, 95% CI for slope = [3.93, 8.17], n= 11) and not for females (Z₉₅,♂=?148.1 +3.11 × (mass♀), r²= 0.58, 95% CI for slope = [‐1.7, 7.9], n= 5, 95% CI for slope= [?1.7, 7.9]). Median depths of presumed foraging dives of harbor seals in the Monterey Bay area were between 5 and 100 m, which were within the range of the previously reported depths for other areas (< 100 m). Our findings generally supported Härkönen's hypothesis that harbor seals forage in the intermediate depth in their environment.     

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.     

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.     

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.     

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.     

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.     

AQUATIC MATING STRATEGIES OF THE MALE PACIFIC HARBOR SEAL (PHOCA VITULINA RICHARDII): ARE MALES DEFENDING THE HOTSPOT?

Compared to the harem and resource defense systems of terrestrial mating pinnipeds, males of aquatic mating species appear unable to monopolize females or resources. We investigated movements, acoustics, and aquatic territorial behavior of male harbor seals, Phoca vitulina richardii , in Elkhorn Slough, California, using VHF telemetry, hydrophones, and acoustic playback experiments. During the mating season 22 males increased time spent in the water and away from haul-out locations, exhibiting activity patterns similar to Atlantic subspecies. Two acoustic display patterns were observed. At one location multiple males aggregated to display with acoustic activity peaking one month before peak estrus. At two other locations, lone males displayed primarily during peak estrus. Acoustic display areas were non-adjacent with a mean ± SE size of 4,228 ± 576 m², similar to harbor seal display patterns in the Moray Firth, Scotland. Underwater playbacks of male vocalizations were used to define territorial boundaries by inducing responses from territory-holding males. Four solitary males defended adjacent territories (mean area 39,571 ± 18,818 m²) along a travel corridor, similar to observations of harbor seals at Miquelon, Newfoundland. Acoustic display stations appeared to be subcomponents of larger territories. Males exhibited site fidelity to territories for at least 2–4 yr. Females moved through territories freely. The establishment of male-display territories along female-traffic corridors resembles terrestrial systems described as hotspot leks.     

A PREHISTORIC BREEDING POPULATION OF HARP SEALS (PHOCA GROENLANDICA) IN THE BALTIC SEA

The pelagic and gregarious, low Arctic harp seal ( Phoca groenlandica ) is the most common seal species in most refuse faunas from coastal hunter-gatherer sites dating from the late Atlantic to the early Subboreal period ( ca. 4000-2000 cal B. C.) in the Baltic Sea. Our main objective was to examine the migration contra breeding population hypotheses regarding the Baltic harp seals. Analyses of epiphyseal fusion data and osteometry of archeological harp seal remains from 25 dwelling-sites suggest that a local breeding population established itself in the early Subboreal period. In the Middle Neolithic the rookery possibly was situated in the Baltic proper, south of Aland and west of Gotland. The mean adult size of the Baltic harp seals decreased, suggesting minimal genetic exchange with the north Atlantic Ocean population. Genetic drift, interspecific competition, and over-hunting by humans are all factors likely to have contributed to the eventual extinction of harp seals in the Baltic Sea.     

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.     

DIGESTIBILITY OF CRUSTACEANS and CAPELIN IN HARP SEALS (PHOCA GROENLANDICA)

Gross energy intake (GEI), apparent digestible efficiency (%DE), and body-mass change were studied both in recently weaned and in subadult harp seals fed capelin ( Mallotus villosus ) and crustaceans ( Parathemisto libellula and Thysanoessa sp.) ad lib. The median %DE of Parathemisto libellula (with a wet-weight energy density (ED) of 3.9 kJ/g) and Thysanoessa sp. (ED: 2.3 kJ/g) was 81% and 83%, respectively, while the median %DE of capelin (ED: 8.6 kJ/g) was 94% and 93% for subadult harp seals and one harp seal pup, respectively. No correlation between age and %DE was obtained (P > 0.05). Despite ad lib. access to the food, the harp seal pups were unable to maintain body mass while on a crustacean diet, and it is therefore suggested that harp seal pups most likely must supplement their crustacean diets with fish to maintain growth.     

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.     

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

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

VARIABILITY IN THE BLUBBER FATTY ACID COMPOSITION OF RINGED SEALS (PHOCA HISPIDA) ACROSS THE CANADIAN ARCTIC

We determined the blubber fatty acid (FA) composition of 281 ringed seals ( Phoca hispida ) across the Canadian Arctic to make inferences about spatial, temporal, and demographic patterns of foraging. Seals were sampled in nine locations between 1992 and 2004. Regional differences in FA signatures were related to the distance between groups, with the greatest similarity occurring among seals sampled in three locations within the Beaufort Sea-Amundsen Gulf. Seals in the western and southeastern portions of Hudson Bay also had similar FA signatures. Discriminant analysis on seventeen FAs classified ringed seals to their correct geographic region with 95% accuracy. Although location accounted for most of the variability in FA signatures, adult and juvenile ringed seals in Frobisher Bay-Labrador Sea and Jones Sound showed significant FA differences, as did male and female ringed seals in Jones Sound and Qaanaaq. Demographic differences were not detected among ringed seals in the Beaufort Sea-Amundsen Gulf or in western Hudson Bay. Seals off the coast of Labrador showed significant seasonal variability in FA signatures. Overall, seasonal, regional, and demographic patterns in FA signatures were consistent with differences in ringed seal diets, as inferred from stomach content and stable isotope analyses.     

白刺珠心组织及其营养功能的超微结构观察

白刺具厚珠心,珠心组织发达。原胚期,根据超微结构特征,珠心组织大体可区分为三层:外围不活跃细胞层、特化细胞层和胚囊外围解体细胞层。珠心细胞解体时,细胞壁首先以微纤丝状态分散,原生质不同程度破坏,并累积脂滴。近胚囊壁降解后,珠心细胞即成为开放细胞,原生质团游离到胚囊周围。胚囊壁在珠孔区和合点区间断,因之,珠心细胞的游离原生质团可直接迁入胚囊内。在胚囊壁和中央细胞质膜之间存在明显间隙区域,其中有大量壁旁体存在,是珠心细胞降解物质进入胚囊的另一重要途径。     

THREE-DIMENSIONAL DIVING BEHAVIORS OF RINGED SEALS (PHOCA HISPIDA)

Dives of five freely diving ringed seals were classified into three-dimentional movement types. Horizontally convoluted dives, defined as dives with angular velocity > 15°/sec, appeared to be foraging or social dives. Simple dives that did not include convoluted movements (angular velocity < 10°/sec) were considered to be exploration dives. Directional dives with nearly linear horizontal travel (horizontal directionality >0.6, on a scale of 0–1) were presumed to be travel dives. Each three-dimensional dive type was observed with similar frequency in dives with two distinct time-depth profiles: V-shaped profiles in which ascent immediately followed descent, and U-shaped profiles in which >7 sec were spent at depth between descent and ascent. The lack of behavioral differences between dives with distinct time-depth profiles suggested that time-depth profiles are not a reliable means of inferring dive behaviors for ringed seals.     

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.     

INFRARED SENSING OF THE UNDER-SNOW LAIRS OF THE RINGED SEAL

Airborne sensing of emitted infrared (IR) was tested as a means of detecting under-snow lairs of the ringed seal ( Phoca hispida ) in study areas in Resolute Passage, N.W.T. Lairs and regularly used large breathing holes could in some cases be detected by emitted IR from altitudes of up to 180 m. Lairs were more easily detected if they had thin roofs, and detectability was higher before sunrise, and under conditions of low wind, low ambient temperature and high cloud cover.     

DO PHOTOPERIOD AND TEMPERATURE INFLUENCE THE MOLT CYCLE OF PHOCA VITULINA IN CAPTIVITY?

Photoperiod variations are the principal environmental signals entraining circannual activity in mammals, which is also believed to be influenced by changes in temperature and nutrition. Control of the molt in phocids has not been extensively studied with experimental manipulations of environmental factors, but substantial work indicates that photoperiod has a primary influence on mammalian breeding and furring mechanisms. Studies were under taken to reestablish the molt cycle in five harbor seals ( Phoca vitulina ) previously kept in an enclosed facility with a limited air/water temperature gradient and artificial photoperiod. The seals were exposed to an extended temperature gradient during the first period of study, and no molting was observed. The second period of study was characterized by a natural occurring photoperiod of 8 h 53 min light (L)/15 h 07 min dark (D) and a maximal 15 h 31 min L/8h 29 min D duration, and molting occurred in all individuals. These results suggest a prevalent influence of photoperiod variation on the harbor seal molt.