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     

Seasonal change in foraging areas and dive depths of breeding king penguins at Heard Island

The seasonal variation in the foraging behaviour of king penguins (Aptenodytes patagonicus) was studied at Heard Island (53°05′S, 73°30′E) during 1992/1993. On seven occasions throughout the breeding cycle, time-depth-light recorders were deployed on breeding adults to record the dive activities and foraging. Foraging locations changed with season: in autumn and spring 1992, adults foraged between 48–52°S and 74–78°E, about 370 km NNE of Heard Island close to the Polar Front. Two penguins tracked in winter travelled 2220 km east of Heard Island (95°E) along the northern ice limit, and 1220 km south of Heard Island to approximately 65°S, respectively. In spring (October), the penguins again foraged further north than during winter. The foraging area utilised in October overlapped the area where the penguins foraged in March/April. The penguins' diving behaviour also varied seasonally: the modal depth of deep dives (>50 m) increased from about 100 m in February to 220 m in October. Mean dive depths increased from 70 ± 52 m in March 1992 to 160 ± 68 m in August 1992. Penguins dived deep (>50 m) only during daylight hours (16 h in February, 9 h in July). Mean dive durations ranged from 2.9 ± 1.1 min in March 1992 to 5.1 ± 1.2 min in August 1992. Associated with changes in foraging location and dive behaviour was a change in diet composition: during summer the penguins ingested mainly myctophid fish (>90%) while in winter the most important diet item was squid. Accepted: 19 October 1998     

Movements and diving of bearded seal (Erignathus barbatus) mothers and pups during lactation and post-weaning

Eleven bearded seals (Erignathus barbatus) were tagged with satellite-linked dive recorders in Kongsfjorden, Svalbard, Norway, in May 1994. These animals included four mother-pup pairs and three single pups. The seals were tracked for 21–258 days. A total of ˜207,000 dives were recorded. Bearded seal mothers showed limited movements during the nursing and moulting periods. After weaning, the pups moved out of the tagging area and dispersed coastally. One pup left Svalbard and moved far offshore to Greenland and Jan Mayen. Bearded seal adults displayed a bi-modal dive behaviour, with peaks of activity that were shallower than 10 m or from 50 to 70 m. Most dives for adult seals (97%) were shorter than 10 min. Young pups performed dives that were shallower and shorter in duration than their accompanying mothers, but diving skills improved rapidly with age. Six of the seven pups dived deeper than 448 m by the time they were 2 months old. Analyses of movement data with respect to separation of mother-pup pairs suggest a lactation period of about 24 days. Accepted: 31 January 2000     

Diving behaviour of hooded seals (Cystophora cristata) in the Greenland and Norwegian Seas

Satellite-linked dive recorders were used to collect data on depths and durations of ∼120,000 dives by 16 hooded seals (Cystophora cristata). Following tagging after moult (four males, eight females) and breeding (four females) off east Greenland, seals dispersed widely in the northeast Atlantic during 172 ± 97 days (mean satellite-linked dive recorder lifetime ± SD). Meso/bathypelagic dives of 5- to 25-min duration to 100–600 m dominated (75%), but some very deep (≥1016 m) and long (>52 min) dives occurred. Diving in open ocean was continuous, with an estimated 90.7±0.8% (mean±SE) of time spent submerged. The proportion of time spent submerged was similar during night and day, but dives during the day were generally deeper and longer (P < 0.05) than during the night. Also, dives in winter were deeper and longer than in summer. Published data on the distribution of likely prey suggest that Greenland halibut (Reinhardtius hippoglossoides), redfish (Sebastes spp.), polar cod (Boreogadus saida), herring (Clupea harengus), squid (Gonatus fabricii) and blue whiting (Micromesistius poutassou) are important prey of hooded seals. Accepted: 22 January 1999     

Distribution and diving of harbour seals (Phoca vitulina) in Svalbard

The distribution, movements and diving of high-arctic harbour seals (Phoca vitulina) were studied in Svalbard, Norway, from 1992 to 1995. A total of 14 seals were equipped with satellite transmitters at Prins Karls Forland (ca. 78°30′N 12°E). These gave data on position, but ten also gave information on dive depths (N ∼ 160,000) and dive durations (N ∼ 162,000). Dive-depth frequencies show that ∼50% of the diving is shallower than 40 m, and that 95% of the diving is shallower than 250 m. Based on dive-duration frequencies, ∼50% of the dives lasted 2–4 min, 90% of the dives lasted less than 7 min, and 97% were shorter than 10 min. All but three seals stayed in the tagging area. Accepted: 6 October 2000     

Satellite tracking and diving behaviour of sub-adult narwhals (<Emphasis Type="Italic">Monodon monoceros</Emphasis>) in Svalbard,Norway

Three juvenile narwhals captured during August 1998 in the northeast of Svalbard, Norway, were equipped with satellite-relayed data loggers (SRDLs) that transmitted diving and swim-speed data, in addition to location, for up to 46 days. A total of 1,354 complete dive cycles were recorded. Most of the diving was shallow and of short duration. Maximum recorded dive depth was 546 m, maximum recorded dive duration was 24.8 min, and maximum recorded swim-speed was 4.7 ms⁻¹. Ascent speed, vertical ascent speed, descent speed and vertical descent speed were all significantly higher during deep dives (>200 m) than for shallow dives (<200 m). In addition both ascent and descent angles were much steeper for deep dives than during shallow dives. Most of the shallow diving seemed to be associated with travelling, with the animal shifting between various locations, while the deep diving (often to the bottom) for extended periods in some specific areas might have been associated with foraging. Even though the sample size in this study is small, the data are the first information available for movements and diving behaviour of narwhals near Svalbard.     

Diving behaviour of dugongs, Dugong dugon

The diving behaviour of 15 dugongs (Dugong dugon) was documented using time-depth recorders (TDRs), which logged a total of 39,507 dives. The TDRs were deployed on dugongs caught at three study sites in northern Australia: Shark Bay, the Gulf of Carpentaria and Shoalwater Bay. The average time for which the dive data were collected per dugong was 10.4±1.1 (S.E.) days. Overall, these dugongs spent 47% of their daily activities within 1.5 m of the sea surface and 72% less than 3 m from the sea surface. Their mean maximum dive depth was 4.8±0.4 m (S.E.), mean dive duration was 2.7±0.17 min and the number of dives per hour averaged 11.8±1.2. The maximum dive depth recorded was 20.5 m; the maximum dive time in water >1.5 m deep was 12.3 min. The effects of dugong sex, location (study site), time of day and tidal cycle on diving rates (dives per hour), mean maximum dive depths, durations of dives, and time spent ≤1.5 m from the surface were investigated using weighted split-plot analysis of variance. The dugongs exhibited substantial interindividual variation in all dive parameters. The interaction between location and time of day was significant for diving rates, mean maximum dive depths and time spent within 1.5 m of the surface. In all these cases, there was substantial variation among individuals within locations among times of day. Thus, it was the variation among individuals that dominated all other effects. Dives were categorised into five types based on the shape of the time-depth profile. Of these, 67% of dives were interpreted as feeding dives (square and U-shaped), 8% as exploratory dives (V-shaped), 22% as travelling dives (shallow-erratic) and 3% as shallow resting dives. There was systematic variation in the distribution of dive types among the factors examined. Most of this variation was among individuals, but this differed across both time of day and tidal state. Not surprisingly, there was a positive relationship between dive duration and depth and a negative relationship between the number of dives per hour and the time spent within 1.5 m of the surface after a dive.     

Diving behavior of immature hawksbill turtles (Eretmochelys imbricata) in a caribbean reef habitat

 Time-depth recorders were deployed on immature hawksbill turtles at the southwestern reefs of Mona Island, Puerto Rico, to examine patterns of diving behavior. Diving profiles of 10–12 day duration were obtained from five turtles ranging in carapace length from 27–52 cm. Turtles exhibited contrasting diurnal and nocturnal diving behaviors. During daylight hours, dives were made 92% of the time, featured continuous depth variation and were attributed to foraging activity. Foraging dive duration increased with turtle size; individual mean dive durations ranged from 19–26 min; mean post-dive surface intervals ranged from 37–64 s; mean depths ranged from 8–10 m. At night, dives were made 86% of the time to constant depths and were interpreted as resting behavior. Resting dive durations were not dependent on turtle size; individual mean dive durations ranged from 35–47 min; mean post-dive surface intervals ranged from 36–60 s; and mean depths from 7–10 m. Immature hawksbill turtles maintained short term home ranges several hundred meters in extension. Accepted: 2 July 1996     

A coral damage index and its application to diving sites in the Egyptian Red Sea

 A coral damage index (CDI) is provided, to screen sites to obtain a perspective on the extent and severity of physical damage to coral. Sites are listed as “hot spots” if in any transect the percent of broken coral colonies (BCC) is greater than or equal to 4% or if the percent cover of coral rubble (CR) is greater than or equal to 3%. To demonstrate its utility, the CDI is applied to a real-life management situation off Hurghada and Safaga, Egypt in the Red Sea. The extent of coral damage covered all four diving sites. Forty percent of all the transects were “hot spots” that required management action. Thirty-one percent of the 16 “hot spot” transects were identified by both broken coral and rubble criteria, 25% by only broken coral criterion and 44% by only coral rubble criterion of the CDI, suggesting that past breakage was responsible for most of the observed damage. Sixty-three percent of the “hot spot” transects were at 4 m depth versus 37% at 8 m depth, suggesting that most of the damage was caused by anchors dragging across the reef in shallow water. The severity of coral damage, reflected by CR, was the greatest at Small Giftun in transect 5 at 4 m depth (333% above the CDI). EI Fanous experienced the most severe degree of broken coral damage (325% above the CDI) at 8 m depth along transect 2. Estimates of the number of dives per year show diving carrying capacities for El Fanous, Gotta Abu Ramada, Ras Abu Soma and Small Giftun being exceeded by large amounts. The CDI can be used globally to; gauge the severity and extent of damage, focus managers on areas that need mooring buoys and associated dive site management programs, and provide a starting point from which to focus more detailed coral reef assessments and restoration programs. Accepted: 30 June 1999     

Three-dimensional dive profiles of free-ranging Weddell seals

We used an acoustic tracking system to record under-ice movements of two free-ranging adult male Weddell seals. The two males were unconstrained and interacting with other Weddell seals at a breeding colony in McMurdo Sound, Antarctica. We reconstructed three-dimensional paths of 279 dives by these seals. All dives were less than 20-min duration and none were deeper than 220 m. These three-dimensional dive profiles were compared with conventional time-depth dive profiles recorded using microprocessor loggers. We assigned each of the 279 dives to 1 of 6 classes using an existing classification scheme on the basis of the time-depth trace. Within these, two-dimensionally derived, classes the actual three-dimensional dive profiles at times varied profoundly. Additional parameters obtained with the acoustic system, such as bearing and distance travelled between diving and surfacing points, demonstrate that significant, additional, biologically important information can be derived from the three-dimensional data. Accepted: 19 December 1999     

Vertical movement of dolphinfish <Emphasis Type="Italic">Coryphaena hippurus</Emphasis> as recorded by acceleration data-loggers in the northern East China Sea

Environmental changes influence foraging behavior for most animals. Dolphinfish, Coryphaena hippurus, are epipelagic predators and have a cosmopolitan tropical to warm-temperate (>20°C) distribution. We simultaneously obtained the ambient temperature and the foraging behavior (i.e., swimming speed, depth and tailbeat acceleration) of dolphinfish, using an acceleration data-logger in May, September, October, November 2007, June 2008, May and July 2010 for 8 individuals. Although the dolphinfish spent a mean ± standard deviation of 43.4 ± 27.7% of their time at the surface (0–5 m), dive excursions from the surface (DES) were observed in all individuals and maximum DES depths ranged from 50.1 to 95.4 m. DES events resulted dives below the thermocline for these dolphinfish, and there was a significantly positive relationship between the isothermal layer depth (ILD) and DES depth. Our results demonstrate that dolphinfish avoided the rapid thermal change beyond the thermocline, and their prey is most likely found in the upper layers of the thermocline. Gliding behavior during the DES phase was also observed and dolphinfish gradually descended to deeper waters with gliding. The gliding time was longer when the ILD was deeper, and fish tended to dive deeper. We suggest that dolphinfish adopt gliding behavior to search a broader range of depths for prey, while minimizing energy use.     

Distribution of abundance, biomass, production and productivity of macrozoobenthos in the sub-Antarctic Magellan Province (South America)

Distribution of abundance, biomass, productivity and production of macrozoobenthos was investigated in four study areas in the Magellan region (South Patagonian Ice-Field, Strait of Magellan, Beagle Channel, Continental Shelf). Using a Reineck box corer and a multibox corer, a total of 277 quantitative benthos samples were taken at 78 stations in water depths between 8 and 1139 m during the Joint Chilean-German-Italian Magellan “Victor-Hensen Campaign” in 1994, the “Polarstern” expedition ANT XIII/4 in 1996 and the Chilean expeditions “Cimar Fiordo II + III” in 1996 and 1997, respectively, on board RV “Vidal Gormaz”. Mean abundance in the South Patagonian Ice-Field was significantly lower than in the Strait of Magellan and the Beagle Channel. Biomass and abundance decreased clearly with depth (20–300 m to 700–1500 m: 3.9 gC m⁻² to 0.6 gC m⁻²; 2832 ind. m⁻² to 569 ind. m⁻²). Average abundance, biomass and production of the whole Magellan region are lower (2318 ind. m⁻², 3.2 gC m⁻², 0.62 gC m⁻² year⁻¹) than in the high Antarctic Weddell Sea. In the Magellan region, macrozoobenthos composition of abundance is mainly dominated by polychaetes (56%), followed by arthropods (16%), echinoderms (10%) and molluscs (11%). Comparisons of our present results with those of high Antarctic areas make it clear that the Magellan region has a transitional character. Accepted: 21 December 1998     

Diving behavior and movements of juvenile hawksbill turtles Eretmochelys imbricata on a Caribbean coral reef

As historically abundant spongivores, hawksbill turtles Eretmochelys imbricata likely played a key ecological role on coral reefs. However, coral reefs are now experiencing global declines and many hawksbill populations are critically reduced. For endangered species, tracking movement has been recognized as fundamental to management. Since movements in marine vertebrates encompass three dimensions, evaluation of diving behavior and range is required to characterize marine turtle habitat. In this study, habitat use of hawksbill turtles on a Caribbean coral reef was elucidated by quantifying diel depth utilization and movements in relation to the boundaries of marine protected areas. Time depth recorders (TDRs) and ultrasonic tags were deployed on 21 Cayman Islands hawksbills, ranging in size from 26.4 to 58.4 cm straight carapace length. Study animals displayed pronounced diel patterns of diurnal activity and nocturnal resting, where diurnal dives were significantly shorter, deeper, and more active. Mean diurnal dive depth (±SD) was 8 ± 5 m, range 2–20 m, mean nocturnal dive depth was 5 ± 5 m, range 1–14 m, and maximum diurnal dive depth was 43 ± 27 m, range 7–91 m. Larger individuals performed significantly longer dives. Body mass was significantly correlated with mean dive depth for nocturnal but not diurnal dives. However, maximum diurnal dive depth was significantly correlated with body mass, suggesting partitioning of vertical habitat by size. Thus, variable dive capacity may reduce intraspecific competition and provide resistance to degradation in shallow habitats. Larger hawksbills may also represent important predators on deep reefs, creating a broad ecological footprint over a range of depths. Communicated by Biology Editor Dr Mark McCormick     

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     

Diving and foraging behaviour of Adélie penguins in areas with and without fast sea-ice

The diving and foraging behaviours of Adélie penguins, Pygoscelis adeliae, rearing chiks at Hukuro Cove, Lützow-Holm Bay, where the fast sea-ice remained throughout summer, were compared to those of penguins at Magnetic Island, Prydz Bay, where the fast sea-ice disappeared in early January. Parent penguins at Hukuro Cove made shallower (7.1–11.3 m) but longer (90–111 s) dives than those at Magnetic Island (22.9 m and 62 s). Dive duration correlated with dive depth at both colonies (r ² = 0.001 ∼ 0.90), but the penguins atg Hukuro Cove made longer dives for a given depth. Parents at Hukuro Cove made shorter foraging trips (8.1–14.4 h) with proportionally longer walking/swimming (diving < 1 m) travel time (27–40% of trip duration) and returned with smaller meals (253–293 g) than those at Magnetic Island, which foraged on average for 57.2 h, spent 2% of time walking/swimming ( < 1 m) travel, and with meals averaging 525 g. Trip duration at both colonies correlated to the total time spent diving. Trip duration at Hukuro Cove, but not at Magnetic Island, increased as walking/swimming ( < 1 m) travel time increased. These differences in foraging behaviour between colonies probably reflected differences in sea-ice cover and the availability of foraging sites. Received: 3 November 1995/Accepted: 29 May 1996     

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.     

Diving pattern and performance in relation to foraging ecology in the gentoo penguin, Pygoscelis papua

We present data on diving pattern and performance (dive depth, duration, frequency and organization during the foraging trip) in gentoo penguins Pygoscelis papua , obtained using time-depth recorders ( n = 9 birds, 99 foraging trips). These data are used to estimate various parameters of foraging activity, e.g. foraging range, prey capture rates, and are compared in relation to breeding chronology. Foraging trip duration was 6 h and 10 h, and trip frequency 1.0/day and 0.96/day, during the brooding and creche periods, respectively. Birds spent on average 52%of each foraging trip diving. Dive depth and duration were highly bimodal: shallow dives (< 21 m) averaged 4 m and 0.23 min, and deep dives (> 30 m) 80 m and 2.5 min, respectively. Birds spent on average 71%and 25%of total diving time in deep and shallow dives, respectively. For deep dives, dive duration exceeded the subsequent surface interval, but shallow dives were followed by surface intervals 2–3 times dive duration. We suggest that most shallow dives are searching/exploratory dives and most deep dives are feeding dives. Deep dives showed clear diel patterns averaging 40 m at dawn and dusk and 80–90 m at midday. Estimated foraging ranges were 2.3 km and 4.1 km during the brood and creche period, respectively. Foraging trip duration increased by 4 h between the brood and creche periods but total time spent in deep dives (i.e. time spent feeding) was the same (3 h). Of 99 foraging trips, 56%consisted of only one dive bout and 44%of 2–4 bouts delimited by extended surface intervals > 10 min. We suggest that this pattern of diving activity reflects variation in spatial distribution of prey rather than the effect of physiological constraints on diving ability.     

Diving patterns of a Ross seal (Ommatophoca rossii ) near the eastern coast of the Antarctic Peninsula

In January 1987 we documented the diving patterns of a female Ross seal (Ommatophoca rossii) in the marginal pack-ice zone near the eastern coast of the Antarctic Peninsula for 2 days using a microprocessor-based time-depth recorder. The seal hauled out during the day and dived continually when in the water at night. Dives averaged 110 m deep and 6.4 min long; the deepest dive was 212 m and the longest 9.8 min. Dives were deepest near twilight and shallowest at night; this pattern suggests that the seal's prey, presumably mid-water squid and fish, may have been making vertical migrations or changing predator-avoidance behavior in response to diel light patterns. The dives of this Ross seal were substantially deeper, on average, than those of crabeater seals (Lobodon carcinophagus), which forage in the same areas on Antarctic krill (Euphausia superba). Received: 15 August 1996 / Accepted: 22 February 1997     

Thermoregulation during swimming and diving in bottlenose dolphins, Tursiops truncatus

Heat transfer from the periphery is an important thermoregulatory response in exercising mammals. However, when marine mammals submerge, peripheral vasoconstriction associated with the dive response may preclude heat dissipation at depth. To determine the effects of exercise and diving on thermoregulation in cetaceans, we measured heat flow and skin temperatures of bottlenose dolphins (Tursiops truncatus) trained to follow a boat and to dive to 15 m. The results demonstrated that skin temperatures usually remained within 1 °C of the water after all exercise levels. Heat flow from peripheral sites (dorsal fin and flukes) increased over resting values immediately after exercise at the water surface and remained elevated for up to 20 min. However, post-exercise values for heat flow from the flukes and dorsal fin decreased by 30–67% when dolphins stationed at 15 m below the surface. The pattern in heat flow was reversed during ascent. For example, mean heat flow from the flukes measured at 5 m depth, 40.10 ± 2.47 W · m⁻², increased by 103.2% upon ascent. There is some flexibility in the balance between thermal and diving responses of dolphins. During high heat loads, heat transfer may momentarily increase during submergence. However, the majority of excess heat in dolphins appears to be dissipated upon resurfacing, thereby preserving the oxygen-conserving benefits of the dive response. Accepted: 4 January 1999     

ANALYSIS OF SWIM VELOCITIES DURING DEEP AND SHALLOW DIVES OF TWO NORTHERN FUR SEALS, CALLORHINUS URSINUS

Swim velocities at 15-sec intervals and maximum depth per dive were recorded by microprocessor units on two "mixed diver" adult female northern fur seals during summer foraging trips. These records allowed comparison of swim velocities of deep (>75 m) and shallow (<75 m) dives.
Deep dives averaged 120 m depth and 3 min duration; shallow dives averaged 30 m and 1.2 min. Mean swim velocities on deep dives were 1.8 and 1.5 m/sec for the two animals; mean swim velocities on shallow dives were 1.5 and 1.2 m/sec. The number of minutes per hour spent diving during the deep and shallow dive patterns were 11 and 27 min, respectively.
Swim velocity, and hence, relative metabolic rate, did not account for the differences in dive durations between deep and shallow dives. The long surface durations associated with deep dives, and estimates of metabolic rates for the observed swim velocities, suggest that deep dives involve significant anaerobic metabolism.