Movements and dive behaviour of two leopard seals (<Emphasis Type="Italic">Hydrurga leptonyx</Emphasis>) off Queen Maud Land,Antarctica

Two adult female leopard seals (Hydrurga leptonyx) were tagged with satellite-linked dive recorders off Queen Maud Land, Antarctica, just after moulting in mid-February. The transmitters transmitted for 80 and 220 days, respectively. Both seals remained within the pack ice relatively close to the Antarctic Continent until early May, when contact was lost with one seal. The one remaining seal then migrated north, to the east side of the South Sandwich Islands in 3 weeks, whereafter it headed east, until contact was lost at 55°S in early September. From mid-May to late September this animal always stayed close to the edge of the pack ice. Both seals made mostly short (<5 min) dives to depths of 10–50 m and only occasionally dove deeper than 200 m, the deepest dive recorded being 304 m. A nocturnal diving pattern was evident in autumn and early winter, while day-time diving prevailed in mid-winter. Haul out probability was highest at mid-day (about 40% in late February and more than 80% in March and April). From May till September the remaining animal mainly stayed at sea, in the vicinity of the pack ice, with only occasional haul outs. These data suggest that a portion of the adult leopard seals may spend the winter mainly in open water, off the edge of the pack ice, where they primarily hunt near the surface. In that case, it is likely that krill (Euphausia superba), as well as penguins, young crabeater seals (Lobodon carcinophaga) and a variety of fish are important prey items.     

Serum chemistry and antibodies against pathogens in antarctic fur seals, Weddell seals, crabeater seals, and Ross seals

Information on health parameters, such as antibody prevalences and serum chemistry that can reveal exposure to pathogens, disease, and abnormal physiologic conditions, is scarce for Antarctic seal species. Serum samples from Antarctic fur seals (Arctocephalus gazella, n=88) from Bouvet?ya (2000-2001 and 2001-2002), and from Weddell seals (Leptonychotes weddellii, n=20), Ross seals (Ommatophoca rossii, n=20), and crabeater seals (Lobodon carcinophagus, n=9) from the pack-ice off Queen Maud Land, Antarctica (2001) were analyzed for enzyme activity, and concentrations of protein, metabolites, minerals, and cortisol. Adult Antarctic fur seal males had elevated levels of total protein (range 64-99 g/l) compared to adult females and pups (range 52-79 g/l). Antarctic fur seals had higher enzyme activities of creatine kinase, lactate dehydrogenase, and amylase, compared to Weddell, Ross, and crabeater seals. Antibodies against Brucella spp. were detected in Weddell seals (37%), Ross seals (5%), and crabeater seals (11%), but not in Antarctic fur seals. Antibodies against phocine herpesvirus 1 were detected in all species examined (Antarctic fur seals, 58%; Weddell seals, 100%; Ross seals, 15%; and crabeater seals, 44%). No antibodies against Trichinella spp., Toxoplasma, or phocine distemper virus (PDV) were detected (Antarctic fur seals were not tested for PDV antibodies). Antarctic seals are challenged by reduced sea ice and increasing temperatures due to climate change, and increased anthropogenic activity can introduce new pathogens to these vulnerable ecosystems and represent a threat for these animals. Our data provide a baseline for future monitoring of health parameters of these Antarctic seal species, for tracking the impact of environmental, climatic, and anthropogenic changes in Antarctica over time.     

Diving and haulout behavior of crabeater seals in the Weddell Sea,Antarctica, during March 1986

Summary Time-depth recorders were used to study the diving and haulout behavior of six crabeater seals in the marginal. ice edge zone of the Weddell Sea during March 1986. Haulout patterns revealed the seals' clear preference for diving during darkness and hauling out onto sea ice during daylight. Seals did not necessarily haul out every day; individual seals hauled out on 80–100% of days during the study period. Four general dive types were identified: 1) traveling dives, 2) foraging dives, 3) crepuscular foraging dives, and 4) exploratory dives. Nearly continual diving occurred for extended periods (about 16 h) nightly, with one individual diving up to 44 h without interruption. Foraging dives occurring during crepuscular periods were deeper than those made during the darkest hours. The authors suggest that the distinct diel pattern of dive timing and depth may be related to possible predator avoidance behavior by the seals' principal prey, Antarctic Krill.     

Ross seal (<Emphasis Type="Italic">Ommatophoca rossii</Emphasis>) annual distribution,diving behaviour,breeding and moulting,off Queen Maud Land,Antarctica

Six out of ten adult Ross seals that were tagged with Argos satellite-linked dive recorders off Queen Maud Land, just after the moult in February, provided data on location and diving activity throughout a year. Shortly after tagging, the animals headed 2,000 km north and stayed pelagic in the area south of the Antarctic Polar Front, until October when they went south into the pack-ice. Throughout the year they made about 100 dives a day, most to a depth of 100–300 m, the deepest dive on record being 792 m, while some dives were very shallow during their stay in the pack-ice. Most dives, outside the breeding and moulting period, lasted for 5–15 min throughout the year. This diving behaviour is consistent with feeding on mid-water fish, like Pleurogramma antarcticum, squid, and to some extent krill (Euphausia superba), when in the pack-ice, and myctophid fish and several species of squid, when in the open ocean. The nursing period was 13 days in mid-November, and moulting occurs in late January–early February, which is the period when sightings surveys for this species should be done.     

CRABEATER SEALS LOBODON CARCINOPHAGUS DURING THE BREEDING SEASON: OBSERVATIONS ON FIVE GROUPS NEAR ENDERBY LAND, ANTARCTICA

Observations on five groups of crabeater seals were conducted between 29 October and 17 November 1985 in the Antarctic pack ice near 66°S 50°E, off Enderby Land. The pups studied were born in the last half of October. Two of them increased in weight at a rate of approximately 4.2kg per day. Pups were weaned in the first half of November at 80–110 kg. The lactation period was 2–3 wk and thus is one of the briefest among pinnipeds. Pups decreased in weight after weaning. The only pup visited after it left the natal floe must have been feeding, as it had lost only 2 kg in a 10–day period during which it moved 13 km. Molt of lanugo appeared to be influenced more by a pup's weight than by whether or not it was weaned.     

Diving physiology and winter foraging behavior of a juvenile leopard seal (Hydrurga leptonyx)

Diving physiology and at-sea behavior of a juvenile leopard seal (Hydrurga leptonyx) were opportunistically measured in the Antarctic Peninsula during winter 2002. Total body oxygen stores were estimated from measures of hematocrit, hemoglobin, myoglobin, and total blood volume and were used to calculate an aerobic dive limit (ADL). Movement patterns and diving behavior were measured by equipping the seal with a Satellite Relay Data Logger that transmitted data from 8–31 August 2002. The seal remained in a focal area, in contrast to crabeater seals tracked simultaneously. The seal displayed short, shallow dives (mean 2.0±1.4 min, 44±48 m) and spent 99.9% of its time within the estimated ADL of 7.4 min. The shallow diving behavior contradicts previous diet research suggesting Antarctic krill (Euphausia superba) is the primary prey of leopard seals during the winter months as krill were found at deeper depths during this period. These measurements of diving and movement of a leopard seal provide valuable preliminary data necessary to develop future research on the at-sea behavior of an apex predator in the Antarctic ecosystem.     

Movements and diving behavior of weaned Weddell seal (Leptonychotes weddellii ) pups

Between 1993 and 1995, the diving behavior and movement patterns of 23 weaned Weddell seal pups (Leptonychotes weddellii) were tracked in the Ross Sea. Antarctica, using satellite-linked time-depth recorders. Regression analyses revealed that for seals of between 8 and 27 weeks old, age was poorly correlated with the dive depth, duration, or frequency. However, changes in dive parameters suggested that Weddell seal pups were attempting to maximize dive time, but the manner in which this was done depended on age and time of day. Movement patterns indicated that most Weddell seal pups left their natal area by the end of February, and traveled north along the Antarctic continent coastline. Several individuals returned to McMurdo Sound, but others were last located more than 400 km from McMurdo. Routes followed suggest that pups can use the pack ice habitat, but prefer to remain closer to the coastline than do adults. Accepted: 21 July 1998     

Pleistocene population expansions of Antarctic seals

We sequenced a portion ( c . 475 bp) of the mitochondrial control region of three species of Antarctic phocid carnivores (Weddell seal, Leptonychotes weddellii , N  = 181; crabeater seal, Lobodon carcinophaga , N  = 143; and Ross seal, Ommatophoca rossii , N  = 41) that live seasonally or permanently in the fast ice and seasonal pack ice of the western Amundsen and Ross seas of western Antarctica. We resolved 251 haplotypes with a haplotype diversity of 0.98 to 0.99. Bayesian estimates of Θ from the program LAMARC ranged from 0.075 for Weddell seals to 0.576 for crabeater seals. We used the values of theta to estimate female effective population sizes ( N_EF ), which were 40 700 to 63 000 for Weddell seals, 44 400 to 97 800 for Ross seals, and 358 500 to 531 900 for crabeater seals. We used mismatch distributions to test for historical population size expansions. Weddell seals and crabeater seals had significant, unimodal mean pairwise difference distributions ( P  = 0.56 and 0.36, respectively), suggesting that their populations expanded suddenly around 731 000 years ago (Weddell seals) and around 1.6 million years ago (crabeater seals). Both of these expansions occurred during times of intensified glaciations and may have been fostered by expanding pack ice habitat.     

Ringed seal (Pusa hispida) seasonal movements,diving, and haul‐out behavior in the Beaufort,Chukchi, and Bering Seas (2011–2017)

Continued Arctic warming and sea‐ice loss will have important implications for the conservation of ringed seals, a highly ice‐dependent species. A better understanding of their spatial ecology will help characterize emerging ecological trends and inform management decisions. We deployed satellite transmitters on ringed seals in the summers of 2011, 2014, and 2016 near Utqia?vik (formerly Barrow), Alaska, to monitor their movements, diving, and haul‐out behavior. We present analyses of tracking and dive data provided by 17 seals that were tracked until at least January of the following year. Seals mostly ranged north of Utqia?vik in the Beaufort and Chukchi Seas during summer before moving into the southern Chukchi and Bering Seas during winter. In all seasons, ringed seals occupied a diversity of habitats and spatial distributions, from near shore and localized, to far offshore and wide‐ranging in drifting sea ice. Continental shelf waters were occupied for >96% of tracking days, during which repetitive diving (suggestive of foraging) primarily to the seafloor was the most frequent activity. From mid‐summer to early fall, 12 seals made ~1‐week forays off‐shelf to the deep Arctic Basin, most reaching the retreating pack‐ice, where they spent most of their time hauled out. Diel activity patterns suggested greater allocation of foraging efforts to midday hours. Haul‐out patterns were complementary, occurring mostly at night until April‐May when midday hours were preferred. Ringed seals captured in 2011—concurrent with an unusual mortality event that affected all ice‐seal species—differed morphologically and behaviorally from seals captured in other years. Speculations about the physiology of molting and its role in energetics, habitat use, and behavior are discussed; along with possible evidence of purported ringed seal ecotypes.     

Diving behaviour and foraging location of female southern elephant seals from Patagonia

Our aim was to describe the free-ranging diving pattern and to determine the location of foraging of pregnant female southern elephant seals, Mirounga leonina , from Peninsula Valdes, Argentina. This colony is unusual in two respects: it is removed from deep water by a broad shallow shelf (345–630 km wide), and colony numbers have been increasing in recent years in contrast to numbers from other southern hemisphere colonies that are stable or in decline. Microprocessor controlled, geolocation-time-depth recorders were deployed on four females, recording a total of 15,836 dives (270 dive days) during the period February to April, 1992. Departing seals crossed the continental shelf quickly (54–5–62–1 h) and did not show signs of foraging until reaching deep water, due east of the colony in the South Atlantic Ocean. Diving was virtually continuous (93% of the time underwater) with overall mean (±S.D.) rates of 2.5±0.2 dives/h, mean dive durations of 22.8 ± 7.1 min (maximum dive duration = 79 min) with 1.6±0.6min surface intervals between dives, and dive depths of 431±193m (maximum dive depth = 1,072 m). The diving pattern of females from Patagonia is similar to that of seals from colonies where numbers are decreasing (Macquarie stock) or are stable (South Georgia Island). Our subjects did not, however, feed in or south of the Antarctic Polar Front, or in cold waters along the Antarctic coast, where seals from declining or stable colonies forage.     

Distribution and abundance of spotted seals Phoca largha and ribbon seals Phoca fasciata in the southern Sea of Okhotsk

The distribution and abundance of spotted seals (Phoca largha) and ribbon seals (Phoca fasciata) were assessed in March and April, 2000, by aerial line-transect surveys along the southern edge of the pack ice off the coast of Hokkaido (southern Sea of Okhotsk), Japan. Five hundred and seventeen spotted seals and 107 ribbon seals were found on the total 2944 km survey line. Total abundance was estimated to be 13 653 spotted (95% CI = 6167–30 252) and 2260 ribbon seals (95% CI = 783–6607) in March, and 6545 spotted (95% CI = 3284–815 644) and 3134 ribbon seals (95% CI = 1247–17 802 512) in April. The pack ice area off Hokkaido had higher densities (0.54 seals km^–2 and 0.58 seals km^–2 in March and April, respectively) of spotted seals than those reported in eastern Sakhalin, whereas densities (0.09 seals km^–2 in March and 0.28 seals km^–2 in April) of ribbon seals were lower than those in eastern Sakhalin. The large number of spotted seal pups suggests that the study area is an important breeding center. A greater number of female spotted seals with pups tended to be found in the center of larger and rougher floes than in other categories, and they were more abundant in stable pack ice areas. Observations of ribbon seals were limited because the survey period preceded the peak of pupping season. Ribbon seal surveys were also hampered by the inability to fly over the main breeding area between the Shiretoko Peninsula and Kunashiri Island.     

Effects of hydrographic variability on the spatial, seasonal and diel diving patterns of southern elephant seals in the eastern Weddell Sea

Weddell Sea hydrography and circulation is driven by influx of Circumpolar Deep Water (CDW) from the Antarctic Circumpolar Current (ACC) at its eastern margin. Entrainment and upwelling of this high-nutrient, oxygen-depleted water mass within the Weddell Gyre also supports the mesopelagic ecosystem within the gyre and the rich benthic community along the Antarctic shelf. We used Conductivity-Temperature-Depth Satellite Relay Data Loggers (CTD-SRDLs) to examine the importance of hydrographic variability, ice cover and season on the movements and diving behavior of southern elephant seals in the eastern Weddell Sea region during their overwinter feeding trips from Bouvetøya. We developed a model describing diving depth as a function of local time of day to account for diel variation in diving behavior. Seals feeding in pelagic ice-free waters during the summer months displayed clear diel variation, with daytime dives reaching 500-1500 m and night-time targeting of the subsurface temperature and salinity maxima characteristic of CDW around 150–300 meters. This pattern was especially clear in the Weddell Cold and Warm Regimes within the gyre, occurred in the ACC, but was absent at the Dronning Maud Land shelf region where seals fed benthically. Diel variation was almost absent in pelagic feeding areas covered by winter sea ice, where seals targeted deep layers around 500–700 meters. Thus, elephant seals appear to switch between feeding strategies when moving between oceanic regimes or in response to seasonal environmental conditions. While they are on the shelf, they exploit the locally-rich benthic ecosystem, while diel patterns in pelagic waters in summer are probably a response to strong vertical migration patterns within the copepod-based pelagic food web. Behavioral flexibility that permits such switching between different feeding strategies may have important consequences regarding the potential for southern elephant seals to adapt to variability or systematic changes in their environment resulting from climate change.     

Distribution and diving behaviour of harp seals (<Emphasis Type="Italic">Pagophilus groenlandicus</Emphasis>) from the Greenland Sea stock

The distribution and diving behaviour of 16 adult harp seals (Pagophilus groenlandicus) from the Greenland Sea stock were studied in 1993 and 1999, using satellite-linked dive recorders (SDRs). The seals remained near the pack-ice edge in the Greenland Sea between breeding and moulting (April/May 1993; 6F) and during the first 7 weeks after moulting (June/July, 1999; 4F, 6M), there diving to depths of <100 m. In mid-July 1999, seven out of eight seals with active SDRs migrated into the Barents Sea, there diving to <400 m and sharing feeding grounds with the Barents Sea harp seal stock. Between September and December, six of these seals joined the eighth seal in the Denmark Strait until March 2000, there diving to depths of 100–400 m. Overall, dives were significantly deeper in the day and in winter than at night and in summer, with some regional differences. Harp seals are considered pack-ice-associated seals, but our tagged seals spent a considerable proportion of their time in open water, their distribution largely overlapping with that of capelin (Mallotus villosus).     

Cooperative hunting behavior,prey selectivity and prey handling by pack ice killer whales (Orcinus orca), type B,in Antarctic Peninsula waters

Currently, there are three recognized ecotypes (or species) of killer whales (Orcinus orca) in Antarctic waters, including type B, a putative prey specialist on seals, which we refer to as “pack ice killer whale” (PI killer whale). During January 2009, we spent a total of 75.4 h observing three different groups of PI killer whales hunting off the western Antarctic Peninsula. Observed prey taken included 16 seals and 1 Antarctic minke whale (Balaenoptera bonaerensis). Weddell seals (Leptonychotes weddellii) were taken almost exclusively (14/15 identified seal kills), despite the fact that they represented only 15% of 365 seals identified on ice floes; the whales entirely avoided taking crabeater seals (Lobodon carcinophaga; 82% relative abundance) and leopard seals (Hydrurga leptonyx; 3%). Of the seals killed, the whales took 12/14 (86%) off ice floes using a cooperative wave‐washing behavior; they produced 120 waves during 22 separate attacks and successfully took 12/16 (75%) of the Weddell seals attacked. The mean number of waves produced per successful attack was 4.1 (range 1–10) and the mean attack duration was 30.4 min (range 15–62). Seal remains that we examined from one of the kills provided evidence of meticulous postmortem prey processing perhaps best termed “butchering.”     

Diving behaviour in relation to water temperature in the southern elephant seal: foraging implications

Summary A time-depth-temperature recorder provided a continuous record of diving by a female southern elephant seal in relation to water temperature for 27 days (1939 dives) after completion of moult. Mean maximum dive depth was 391±2.6 m and the overall maximum was 775 m. Dives lasted on average 17.5±0.09 min. Most dives showed a rapid descent to the discontinuity between the cold surface water and warmer deep water. Consequently the seal spent 57% of its time while diving at a depth of 200–400 m when it may have been foraging. This strongly suggests that the seal was exploiting a food source at the discontinuity between vertically stratified water masses. The water temperature data also indicated that the seal was diving in waters south of the Antarctic Polar Front and at some distance from the northern edge of the pack ice. The seal spent 88% of its time under water. Normal surface intervals between dives lasted an average of 2.1 ± 0.1 min whereas 16 extended surface intervals (>10 min duration) lasted 32.7±4.6 min. Dives were deeper during the day than at night and all but one extended surface interval occurred at night. The pattern of dives was similar to records from northern elephant seals but this is the first study to show how diving behaviour relates to water temperature.     

Annual distribution of hooded seals (Cystophora cristata) in the Greenland and Norwegian seas

 Nineteen hooded seals (Cystophora cristata) were tagged with satellite-linked platform terminal transmitters (PTT) on the sea ice near Jan Mayen. Fifteen were instrumented after completion of the moult in July 1992 (five males, ten females, at 71°N, 12°W), and four during breeding in March 1993 (four females, at 69°N, 20°W). Sixteen of the seals were tagged with Satellite-Linked Time-Depth-Recorders (SLTDR), yielding location, dive depth and dive duration data. The average (±SD) longevity of all PTTs was 199±84 days (n=19; range: 43–340 days), and they yielded 12,834 location fixes. Between tagging in July 1992 and pupping in March 1993, two seals remained in or near the ice off the east coast of Greenland for most of the tracking period. However, most of the seals made one or several trips away from the ice edge, mostly to distant waters. These excursions had an average (±SD) duration of 47±22 days (n=46; range: 4–99 days). Eight seals travelled to waters off the Faeroe Islands, three to the continental shelf break south of Bear Island, and three to the Irminger Sea southwest of Iceland. Eleven seals were tracked in the period between breeding (March/April) and moulting (July). Several of these spent extended periods at sea west of the British Isles, or in the Norwegian Sea. Received: 3 August 1994/Accepted: 4 July 1995     

Movements and diving of adult ringed seals (Phoca hispida) in Svalbard

Seven post-moulting adult ringed seals (Phoca hispida) were equipped with Satellite Linked Dive Recorders in Svalbard in July 1996 to determine if ringed seals conduct long-distance post-moulting feeding excursions, and to obtain details of their diving behaviour. The mean duration of tags was 206 days (range 103–325). Two seals swam 400 km north to the drifting pack ice (82°N). The rest undertook more local movements. Forty-eight percent of all dives were shallower than 20 m and 90% were shallower than 100 m. Ninety-five percent of all dive durations were shorter than 10 min, and 99.5% were shorter than 15 min. This study has shown that adult ringed seals undertake varying patterns of post-moulting excursions. Accepted: 1 April 2000     

Seasonal diving behaviour in lactating subantarctic fur seals on Amsterdam Island

Diving behaviour was investigated in female subantarctic fur seals (Arctocephalus tropicalis) breeding on Amsterdam Island, Indian Ocean. Data were collected using electronic Time Depth Recorders on 19 seals during their first foraging trip after parturition in December, foraging trips later in summer, and during winter. Subantarctic fur seals at Amsterdam Island are nocturnal, shallow divers. Ninety-nine percent of recorded dives occurred at night. The diel dive pattern and changes in dive parameters throughout the night suggest that fur seals follow the nycthemeral migrations of their main prey. Seasonal changes in diving behaviour amounted to the fur seals performing progressively deeper and longer dives from their first foraging trip through winter. Dive depth and dive duration increased from the first trip after parturition (16.6 ± 0.5 m and 62.1 ± 1.6 s respectively, n=1000) to summer (19.0 ± 0.4 m and 65 ± 1 s, respectively, n=2000) through winter (29.0 ± 1.0 m and 91.2 ± 2.2 s, respectively, n=800). In summer, subantarctic fur seals increased the proportion of time spent at the bottom during dives of between 10 and 20 m, apparently searching for prey when descending to these depths, which corresponded to the oceanic mixed layer. In winter, fur seals behaved similarly when diving between 20 and 50 m, suggesting that the most profitable depths for feeding moved down during the study period. Most of the dives did not exceed the physiological limits of individuals. Although dive frequency did not vary (10 dives/h of night), the vertical travel distance and the time spent diving increased throughout the study period, while the post-dive interval decreased, indicating that subantarctic fur seals showed a greater diving effort in winter, compared to earlier seasons. Accepted: 1 August 1999     

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.     

Diet and prey consumption of Antarctic petrels Thalassoica antarctica at Svarthamaren, Dronning Maud Land, and at sea outside the colony

The diet of the Antarctic petrel Thalassoica antarctica was studied during two seasons at Svarthamaren, an inland colony in Dronning Maud Land, Antarctica, and in the pack ice off the coast of Svarthamaren. The most important food (wet mass) at Svarthamaren was crustaceans (67%), fish (29%) and squid (5%); however, individuals collected in the pack ice took mostly fish (87%). The prey composition and lengths of prey are comparable to what has been documented in other studies on this species. Estimates of food consumption by birds breeding at Svarthamaren (ca. 250,000 pairs) suggest that approximately 6500 tonnes of crustaceans, 2800 tonnes of fish and 435 tonnes of squid are consumed during the breeding season. The annual consumptions of these birds are estimated to be 34,100 tonnes of crustaceans, 14,700 tonnes of fish, and 2300 tonnes of squid. Satellite telemetry data indicate that Antarctic petrels from Svarthamaren may fly more than 3000 km during one foraging trip, and thus may cover a huge ocean area to obtain their prey. Received: 1 September 1997 / Accepted: 3 February 1998