Diving behaviour and energetics in breeding little penguins (Eudyptula minor)

We present data on the diving behaviour and the energetics of breeding little penguins in Tasmania, Australia. Using an 18 m long still water canal in conjunction with respirometry, we determined the energy requirements while diving. Using electronic devices measuring dive depth or swimming speed, we investigated the foraging behaviour at sea. Cost of Transport was calculated to be minimal at the speed the birds prefer at sea (1.8 m/s) and averaged 11.1 J/kg/m (power requirements at that speed: 20.0 W/kg). Metabolic rate of little penguins resting in water was found to be 8.5 W/kg. The externally-attached devices had no significant influence on the energy expenditure.
Foraging trips can be divided into four distinct phases with different diving behaviours. A mean of 500 dives was executed per foraging trip lasting about 18 hours with 60% of this time being spent swimming. The total distance travelled averaged 73 km per day, although foraging range was about 12km. Mean swimming speed of little penguins at sea was 1.8 m/s, maximum swimming speed was 3.3 m/s. More than 50% of all dives had maxima not exceeding 2 m. Maximum depth reached was 27 m. Mean dive duration was 21 s. There were inter-sex differences in diving behaviour as well as changes in foraging behaviour over the breeding period. Aerobic dive limits (ADL) in the wild were estimated between 42 and 50 s. From the swim canal experiments we derived an ADL of 44 s. Total oxygen stores were calculated to be 45 ml O₂/kg. Only 2% of all dives exceeded the ADL. FMRs at sea were calculated to be between 1280 and 1500 kJ/kg/d according to chick size. The yearly food requirements of a breeding little penguin amount to 114 kg.     

Diving behaviour, dive cycles and aerobic dive limit in the platypus Ornithorhynchus anatinus

We investigated the diving behaviour, the time allocation of the dive cycle and the behavioural aerobic dive limit (ADL) of platypuses (Ornithorhynchus anatinus) living at a sub-alpine Tasmanian lake. Individual platypuses were equipped with combined data logger-transmitter packages measuring dive depth. Mean dive duration was 31.3 s with 72% of all dives lasting between 18 and 40 s. Mean surface duration was 10.1 s. Mean dive depth was 1.28 m with a maximum of 8.77 m. Platypuses performed up to 1600 dives per foraging trip with a mean of 75 dives per hour. ADL was estimated by consideration of post-dive surface intervals vs. dive durations. Only 15% of all dives were found to exceed the estimated ADL of 40 s, indicating mainly aerobic diving in the species. Foraging platypuses followed a model of optimised recovery time, the optimal breathing theory. Total bottom duration or total foraging duration per day is proposed as a useful indicator of foraging efficiency and hence habitat quality in the species.     

Effects of diving and swimming behavior on body temperatures of pacific leatherback turtles in tropical seas

Mathematical models and recordings of cloacal temperature suggest that leatherback turtles (Dermochelys coriacea) maintain core body temperature higher than ambient water temperature (T(W)) while freely swimming at sea. We investigated the thermoregulatory capabilities of free-ranging leatherbacks and, specifically, the effect that changes in diving patterns and ambient temperatures have on leatherback body temperatures (T(B)). Data loggers were used to record subcarapace and gastrointestinal tract temperatures (T(SC) and T(GT), respectively), T(W), swim speed, dive depth, and dive times of female leatherback turtles during internesting intervals off the coast of Guanacaste, Costa Rica. Mean T(SC) (28.7 degrees -29.0 degrees C) was significantly higher than mean T(W) (25.0 degrees -27.5 degrees C). There was a significant positive relationship between T(SC) and T(W) and a significant negative correlation between T(SC) and dive depth and T(GT) and dive depth. Rapid fluctuations in T(GT) occurred during the first several days of the internesting interval, which suggests that turtles were ingesting prey or water during this time. Turtles spent 79%-91% of the time at sea swimming at speeds greater than 0.2 m s(-1), and the average swim speed was 0.7 +/- 0.2 m s(-1). Results from this study show that alterations in diving behavior and T(W) affect T(B) of leatherback turtles in the tropics. Body temperatures of free-ranging leatherback turtles correspond well with values for T(B) predicted by mathematical models for tropical conditions.     

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     

Energetic costs of surface swimming and diving of birds

The energetic costs of swimming at the surface (swimming) and swimming underwater (diving) are compared in tufted ducks (Aythya fuligula) and three species of penguins, the gentoo (Pygoscelis papua), the king (Aptenodytes patagonicus), and the emperor (Aythya forsteri). Ducks swim on the surface and use their webbed feet as paddles, whereas penguins tend to swim just below the surface and use their flippers as hydrofoils, the latter being much more efficient. Penguins are more streamlined in shape. Thus, the amount of energy required to transport a given mass of bird a given distance (known as the cost of transport) is some two to three times greater in ducks than in penguins. Ducks are also very buoyant, and overcoming the force of buoyancy accounts for 60% and 85% of the cost of descent and remaining on the bottom, respectively, in these birds. The energy cost of a tufted duck diving to about 1.7 m is similar to that when it is swimming at its maximum sustainable speed at the surface (i.e., approximately 3.5 times the value when resting on water). Nonetheless, because of the relatively short duration of its dives, the tufted duck dives well within its calculated aerobic dive limit (cADL, usable O(2) stores per rate of O(2) usage when underwater). However, these three species of penguins have maximum dive durations ranging from 5 min to almost 16 min and maximum dive depths from 155 to 530 m. When these birds dive, they have to metabolise at no more than when resting in water in order for cADL to encompass the duration of most of their natural dives. In gentoo and king penguins, there is a fall in abdominal temperature during bouts of diving; this may reduce the oxygen requirements in the abdominal region, thus enabling dive duration to be extended further than would otherwise be the case.     

Locomotion in diving elephant seals: physical and physiological constraints

To better understand how elephant seals (Mirounga angustirostris) use negative buoyancy to reduce energy metabolism and prolong dive duration, we modelled the energetic cost of transit and deep foraging dives in an elephant seal. A numerical integration technique was used to model the effects of swim speed, descent and ascent angles, and modes of locomotion (i.e. stroking and gliding) on diving metabolic rate, aerobic dive limit, vertical displacement (maximum dive depth) and horizontal displacement (maximum horizontal distance along a straight line between the beginning and end locations of the dive) for aerobic transit and foraging dives. Realistic values of the various parameters were taken from previous experimental data. Our results indicate that there is little energetic advantage to transit dives with gliding descent compared with horizontal swimming beneath the surface. Other factors such as feeding and predator avoidance may favour diving to depth during migration. Gliding descent showed variable energy savings for foraging dives. Deep mid-water foraging dives showed the greatest energy savings (approx. 18%) as a result of gliding during descent. In contrast, flat-bottom foraging dives with horizontal swimming at a depth of 400m showed less of an energetic advantage with gliding descent, primarily because more of the dive involved stroking. Additional data are needed before the advantages of gliding descent can be fully understood for male and female elephant seals of different age and body composition. This type of data will require animal-borne instruments that can record the behaviour, three-dimensional movements and locomotory performance of free-ranging animals at depth.     

Diving behaviour of a reptile (Crocodylus johnstoni) in the wild: interactions with heart rate and body temperature

The differences in physical properties of air and water pose unique behavioural and physiological demands on semiaquatic animals. The aim of this study was to describe the diving behaviour of the freshwater crocodile Crocodylus johnstoni in the wild and to assess the relationships between diving, body temperature, and heart rate. Time-depth recorders, temperature-sensitive radio transmitters, and heart rate transmitters were deployed on each of six C. johnstoni (4.0-26.5 kg), and data were obtained from five animals. Crocodiles showed the greatest diving activity in the morning (0600-1200 hours) and were least active at night, remaining at the water surface. Surprisingly, activity pattern was asynchronous with thermoregulation, and activity was correlated to light rather than to body temperature. Nonetheless, crocodiles thermoregulated and showed a typical heart rate hysteresis pattern (heart rate during heating greater than heart rate during cooling) in response to heating and cooling. Additionally, dive length decreased with increasing body temperature. Maximum diving length was 119.6 min, but the greatest proportion of diving time was spent on relatively short (<45 min) and shallow (<0.4 m) dives. A bradycardia was observed during diving, although heart rate during submergence was only 12% lower than when animals were at the surface.     

Deep-diving foraging behaviour of sperm whales (Physeter macrocephalus)

1. Digital tags were used to describe diving and vocal behaviour of sperm whales during 198 complete and partial foraging dives made by 37 individual sperm whales in the Atlantic Ocean, the Gulf of Mexico and the Ligurian Sea. 2. The maximum depth of dive averaged by individual differed across the three regions and was 985 m (SD = 124.3), 644 m (123.4) and 827 m (60.3), respectively. An average dive cycle consisted of a 45 min (6.3) dive with a 9 min (3.0) surface interval, with no significant differences among regions. On average, whales spent greater than 72% of their time in foraging dive cycles. 3. Whales produced regular clicks for 81% (4.1) of a dive and 64% (14.6) of the descent phase. The occurrence of buzz vocalizations (also called 'creaks') as an indicator of the foraging phase of a dive showed no difference in mean prey capture attempts per dive between regions [18 buzzes/dive (7.6)]. Sperm whales descended a mean of 392 m (144) from the start of regular clicking to the first buzz, which supports the hypothesis that regular clicks function as a long-range biosonar. 4. There were no significant differences in the duration of the foraging phase [28 min (6.0)] or percentage of the dive duration in the foraging phase [62% (7.3)] between the three regions, with an overall average proportion of time spent actively encountering prey during dive cycles of 0.53 (0.05). Whales maintained their time in the foraging phase by decreasing transit time for deeper foraging dives. 5. Similarity in foraging behaviour in the three regions and high diving efficiencies suggest that the success of sperm whales as mesopelagic predators is due in part to long-range echolocation of deep prey patches, efficient locomotion and a large aerobic capacity during diving.     

Effects of air and water temperatures on resting metabolism of auklets and other diving birds

For small aquatic endotherms, heat loss while floating on water can be a dominant energy cost, and requires accurate estimation in energetics models for different species. We measured resting metabolic rate (RMR) in air and on water for a small diving bird, the Cassin's auklet (Ptychoramphus aleuticus), and compared these results to published data for other diving birds of diverse taxa and sizes. For 8 Cassin's auklets (~165 g), the lower critical temperature was higher on water (21 °C) than in air (16 °C). Lowest values of RMR (W kg?1) averaged 19% higher on water (12.14 ± 3.14 SD) than in air (10.22 ± 1.43). At lower temperatures, RMR averaged 25% higher on water than in air, increasing with similar slope. RMR was higher on water than in air for alcids, cormorants, and small penguins but not for diving ducks, which appear exceptionally resistant to heat loss in water. Changes in RMR (W) with body mass either in air or on water were mostly linear over the 5- to 20-fold body mass ranges of alcids, diving ducks, and penguins, while cormorants showed no relationship of RMR with mass. The often large energetic effects of time spent floating on water can differ substantially among major taxa of diving birds, so that relevant estimates are critical to understanding their patterns of daily energy use.     

The diving behavior of blue and fin whales: is dive duration shorter than expected based on oxygen stores?

Many diving seabirds and marine mammals have been found to regularly exceed their theoretical aerobic dive limit (TADL). No animals have been found to dive for durations that are consistently shorter than their TADL. We attached time-depth recorders to 7 blue whales and 15 fin whales (family Balaenopteridae). The diving behavior of both species was similar, and we distinguished between foraging and traveling dives. Foraging dives in both species were deeper, longer in duration and distinguished by a series of vertical excursions where lunge feeding presumably occurred. Foraging blue whales lunged 2.4 (+/-1.13) times per dive, with a maximum of six times and average vertical excursion of 30.2 (+/-10.04) m. Foraging fin whales lunged 1.7 (+/-0.88) times per dive, with a maximum of eight times and average vertical excursion of 21.2 (+/-4.35) m. The maximum rate of ascent of lunges was higher than the maximum rate of descent in both species, indicating that feeding lunges occurred on ascent. Foraging dives were deeper and longer than non-feeding dives in both species. On average, blue whales dived to 140.0 (+/-46.01) m and 7.8 (+/-1.89) min when foraging, and 67.6 (+/-51.46) m and 4.9 (+/-2.53) min when not foraging. Fin whales dived to 97.9 (+/-32.59) m and 6.3 (+/-1.53) min when foraging and to 59.3 (+/-29.67) m and 4.2 (+/-1.67) min when not foraging. The longest dives recorded for both species, 14.7 min for blue whales and 16.9 min for fin whales, were considerably shorter than the TADL of 31.2 and 28.6 min, respectively. An allometric comparison of seven families diving to an average depth of 80-150 m showed a significant relationship between body mass and dive duration once Balaenopteridae whales, with a mean dive duration of 6.8 min, were excluded from the analysis. Thus, the short dive durations of blue whales and fin whales cannot be explained by the shallow distribution of their prey. We propose instead that short duration diving in large whales results from either: (1) dispersal behavior of prey; or (2) a high energetic cost of foraging.     

SUMMER DIVING BEHAVIOR OF MALE WALRUSES IN BRISTOL BAY, ALASKA

Pacific walruses ( Odobenus rosmarus divergens ) make trips from ice or land haul-out sites to forage for benthic prey. We describe dive and trip characteristics from time-depth-recorder data collected over a one-month period during summer from four male Pacific walruses in Bristol Bay, Alaska. Dives were classified into four types. Shallow (4 m), short (2.7 min), square-shaped dives accounted for 11% of trip time, and many were probably associated with traveling. Shallow (2 m) and very short (0.5 min) dives composed only 1% of trip time. Deep (41 m), long (7.2 min), square-shaped dives accounted for 46% of trip time and were undoubtedly associated with benthic foraging. V-shaped dives ranged widely in depth, were of moderate duration (4.7 min), and composed 3% of trip time. These dives may have been associated with navigation or exploration of the seafloor for potential prey habitat. Surface intervals between dives were similar among dive types, and generally lasted 1–2 min. Total foraging time was strongly correlated with trip duration and there was no apparent diel pattern of diving in any dive type among animals. We found no correlation between dive duration and postdive surface interval within dive types, suggesting that diving occurred within aerobic dive limits. Trip duration varied considerably within and among walruses (0.3–9.4 d), and there was evidence that some of the very short trips were unrelated to foraging. Overall, walruses were in the water for 76.6% of the time, of which 60.3% was spent diving.     

Foraging energetics of arctic cormorants and the evolution of diving birds

Efficient body insulation is assumed to have enabled birds and mammals to colonize polar aquatic ecosystems. We challenge this concept by comparing the bioenergetics of cormorants ( Phalacrocorax carbo ) living in temperate and arctic conditions. We show that although these birds have limited insulation, they maintain high body temperature (42.3 °C) when diving in cold water (1–10 °C). Their energy demand at these times is extremely high (up to 60 W kg⁻¹). Free-living cormorants wintering in Greenland (water temperature −1 °C) profoundly alter their foraging activity, thus minimizing time spent in water and the associated high thermoregulatory costs. They then meet their daily food demand within a single intense dive bout (lasting 9 min on average). Their substantial energy requirements are balanced by the highest predatory efficiency so far recorded for aquatic predators. We postulate that similar behavioural patterns allowed early diving birds (Cretaceous) to colonize cold coastal areas before they evolved efficient insulation.     

Activity Time Budget during Foraging Trips of Emperor Penguins

We developed an automated method using depth and one axis of body acceleration data recorded by animal-borne data loggers to identify activities of penguins over long-term deployments. Using this technique, we evaluated the activity time budget of emperor penguins (n = 10) both in water and on sea ice during foraging trips in chick-rearing season. During the foraging trips, emperor penguins alternated dive bouts (4.8±4.5 h) and rest periods on sea ice (2.5±2.3 h). After recorder deployment and release near the colony, the birds spent 17.9±8.4% of their time traveling until they reached the ice edge. Once at the ice edge, they stayed there more than 4 hours before the first dive. After the first dive, the mean proportions of time spent on the ice and in water were 30.8±7.4% and 69.2±7.4%, respectively. When in the water, they spent 67.9±3.1% of time making dives deeper than 5 m. Dive activity had no typical diurnal pattern for individual birds. While in the water between dives, the birds had short resting periods (1.2±1.7 min) and periods of swimming at depths shallower than 5 m (0.25±0.38 min). When the birds were on the ice, they primarily used time for resting (90.3±4.1% of time) and spent only 9.7±4.1% of time traveling. Thus, it appears that, during foraging trips at sea, emperor penguins traveled during dives >5 m depth, and that sea ice was primarily used for resting. Sea ice probably provides refuge from natural predators such as leopard seals. We also suggest that 24 hours of sunlight and the cycling of dive bouts with short rest periods on sea ice allow emperor penguins to dive continuously throughout the day during foraging trips to sea.     

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.     

The influence of depth on a breath-hold diver: Predicting the diving metabolism of Steller sea lions (Eumetopias jubatus)

Diving animals must endeavor to increase their dive depths and prolong the time they spend exploiting resources at depth. Results from captive and wild studies suggest that many diving animals extend their foraging bouts by decreasing their metabolisms while submerged. We measured metabolic rates of Steller sea lions (Eumetopias jubatus) trained to dive to depth in the open ocean to investigate the relationships between diving behaviour and the energetic costs of diving. We also constructed a general linear model to predict the oxygen consumption of sea lions diving in the wild. The resultant model suggests that swimming distance and depth of dives significantly influence the oxygen consumption of diving Steller sea lions. The predictive power of the model was tested using a cross-validation approach, whereby models reconstructed using data from pairs of sea lions were found to accurately predict the oxygen consumption of the third diving animal. Predicted oxygen consumption during dives to depth ranged from 3.37 L min^− 1 at 10 m, to 1.40 L min^− 1 at 300 m over a standardized swimming distance of 600 m. This equated to an estimated metabolic rate of 97.54 and 40.52 MJ day^− 1, and an estimated daily feeding requirement of 18.92 and 7.96 kg day^− 1 for dives between 10 and 300 m, respectively. The model thereby provides information on the potential energetic consequences that alterations in foraging strategies due to changes in prey availability could have on wild populations of sea lions.     

Recording the free-living behaviour of small-bodied, shallow-diving animals with data loggers

1. Time-depth data recorders (TDRs) have been widely used to explore the behaviour of relatively large, deep divers. However, little is known about the dive behaviour of small, shallow divers such as semi-aquatic mammals. 2. We used high-resolution TDRs to record the diving behaviour of American mink Mustela vison (weight of individuals 580-1275 g) in rivers in Oxfordshire (UK) between December 2005 and March 2006. 3. Dives to > 0.2 m were measured in all individuals (n = 6). Modal dive depth and duration were 0.3 m and 10 s, respectively, although dives up to 3 m and 60 s in duration were recorded. Dive duration increased with dive depth. 4. Temperature data recorded by TDRs covaried with diving behaviour: they were relatively cold (modal temperature 4-6 degrees C across individuals) when mink were diving and relatively warm (modal temperature 24-36 degrees C across individuals) when mink were not diving. 5. Individuals differed hugely in their use of rivers, reflecting foraging plasticity across both terrestrial and aquatic environments. For some individuals there was < 1 dive per day while for others there was > 100 dives per day. 6. We have shown it is now possible to record the diving behaviour of small free-living animals that only dive a few tens of centimetres, opening up the way for a new range of TDR studies on shallow diving species.     

Body temperature and insulation in diving Great Cormorants and European Shags

1. Cormorants are typically considered as wettable diving birds with high thermoregulatory costs and are presumed to exert substantial predatory pressure on fish stocks.
2. The stomach temperatures of seven Great Cormorants and three European Shags were recorded during a total of 108 foraging trips undertaken near the Chausey Islands breeding colony (France).
3. Both species kept a constant body temperature during the dive series which lasted up to 158 min and were conducted in 12°C water. Consequently, assuming that heat loss to the water is equal to heat production in diving Great Cormorants, the minimal insulating plumage air volume was calculated to be 0·371 × 10^–3 m³ (corresponding to a 1·62-mm air layer) in males and 0·347 × 10^–3 m³ (corresponding to a 1·90-mm air layer) in females.
4. Furthermore, it is shown that plumage air volume and dive depth are the major factors influencing heat flux to the water and that the energetics of diving Great Cormorants may also vary substantially according to fat layer thickness, water temperature and body temperature. Swim speed plays only a minor role.
5. Considering these results, it is postulated that Great Cormorants may have optimized plumage air volume so as to minimize both mechanical costs (upthrust) and thermoregulatory costs of swimming in cold, shallow water.
6. Finally, body temperature patterns recorded in different cormorant species while diving are compared.     

Movements, diving and foraging behaviour of grey seals (Halichoerus grypus)

This study presents the first direct observations of the movements and behaviour of free-ranging grey seals at sea. Radio and ultrasonic transmitters were attached to three sub-adult male grey seals which were then tracked from a suitable vessel. Behaviour at sea fell into one of three categories: travelling between haul-outs, short duration trips, and resting adjacent to haul-out sites. Travelling was characterized by direct, relatively fast horizontal movement and by V-shaped dive profiles. During short duration trips the seals swam slower and invariably exhibited square-wave dive profiles, spending approximately 60% of total dive duration at the maximum depth. Resting involved shallow dives close to haul-out sites and an absence of directed lateral movement.
The excellent navigational abilities of grey seals are illustrated by the rapid, direct swimming between distant haul-out sites. It is proposed that short duration trips are specifically for foraging because of their association with other piscivores, and because swimming was slow and mostly on or near the sea bed (grey seals are known to feed almost exclusively on demersal and benthic fish). These trips accounted for only 14% of the nine days that seal 1 was tracked. It is also proposed that the habit of diving to the sea bed whilst travelling between distant haul-out sites is to allow opportunistic foraging with only a small increase in total swimming distance.     

Underwater observations of foraging free-living Atlantic walruses (Odobenus rosmarus rosmarus) and estimates of their food consumption

Food consumption of Atlantic walruses (Odobenus rosmarus rosmarus L.) was quantified by combining underwater observations of feeding with satellite-telemetry data on movement and diving activity. The study was conducted between 31 July and 7 August 2001 in Young Sound (74°N-20°W) in Northeast Greenland. On ten occasions, divers were able to accompany foraging walruses to the sea floor and collect the shells of newly predated bivalves (Mya truncata, Hiatella arctica, Serripes groenlandicus) for determination of number of prey and biomass ingested per dive. Simultaneously, the activity of a 1,200-kg adult male walrus was studied by use of satellite-telemetry during an entire foraging cycle that included 74 h at sea followed by a 23-h rest on land. An average of 53.2 bivalves (SE=5.2, range: 34-89, n=10) were consumed per dive, corresponding to 149.0 g shell-free dry matter (SE=18.9, range: 62.4-253.1 g), or 2,576 kJ per dive (SE=325.2, range: 1,072-4,377 kJ). During the foraging trip, the walrus spent 57% of the time diving to depths of between 6 and 32 m, and it made a total of 412 dives that lasted between 5 and 7 min (i.e. typical foraging dives). If the entire feeding cycle is considered (97 h), the estimated daily gross energy intake was 214 kJ per kg body mass (95% CI: 153-275 kJ), corresponding to the ingestion of 57 kg (95% CI: 41-72 kg) wet weight bivalve biomass per day, or 4.7 (95% CI: 3.3-5.9%) of total walrus body mass. Due to ice cover, walrus access to the plentiful inshore bivalve banks in the area is restricted to the short summer period, where walruses rely on them for replenishing energy stores. It is hypothesised that the documented decrease in the extent and duration of Arctic sea ice may increase food availability for walruses in eastern Greenland in the future.     

Diving behaviour of the Shy Albatross Diomedea cauta in Tasmania: initial findings and dive recorder assessment

The diving behaviour of the Shy Albatross Diomedea cauta was investigated using archival time-depth recorders (TDRs) and maximum depth gauges (MDGs). Data from birds carrying multiple devices and from diving simulations indicated that the degree of correspondence between TDRs and MDGs varied with the dive depth, duration and frequency, as well as with body placement. The MDGs were the most reliable when the diving depth was greater than 0.5 m, when the diving frequency was low and when gauges were placed on the birds' backs. The TDRs were used during late incubation and early chick rearing in 1994. Fifty-two dives (0.4 m) were recorded during 20 foraging trips of 15 individuals. The majority of dives were within the upper 3 m of the water column and lasted for less than 6 s. However, dives to 7.4 m and others lasting 19 s were recorded. The albatrosses dived between 07.00 h and 22.00 h, with peaks in their diving activity near midday and twilight. Mean diving depth varied throughout the day. with the deepest dives occurring between 10.00 h and 12.00 h. Two dive types were identified on the basis of the relationship between dive depth and descent rate. Plunge dives were short (5 s), and the birds reached a maximum depth of 2.9 m. Swimming dives were both longer and deeper. The characteristics of Shy Albatross plunge dives were similar to those of gannets Morus spp., which are known to be proficient plunge divers. Swimming dives suggest that Shy Albatrosses actively pursue prey underwater.