The marine mammal dive response is exercise modulated to maximize aerobic dive duration

When aquatically adapted mammals and birds swim submerged, they exhibit a dive response in which breathing ceases, heart rate slows, and blood flow to peripheral tissues and organs is reduced. The most intense dive response occurs during forced submersion which conserves blood oxygen for the brain and heart, thereby preventing asphyxiation. In free-diving animals, the dive response is less profound, and energy metabolism remains aerobic. However, even this relatively moderate bradycardia seems diametrically opposed to the normal cardiovascular response (i.e., tachycardia and peripheral vasodilation) during physical exertion. As a result, there has been a long-standing paradox regarding how aquatic mammals and birds exercise while submerged. We hypothesized based on cardiovascular modeling that heart rate must increase to ensure adequate oxygen delivery to active muscles. Here, we show that heart rate (HR) does indeed increase with flipper or fluke stroke frequency (SF) during voluntary, aerobic dives in Weddell seals (HR?=?1.48SF?-?8.87) and bottlenose dolphins (HR?=?0.99SF?+?2.46), respectively, two marine mammal species with different evolutionary lineages. These results support our hypothesis that marine mammals maintain aerobic muscle metabolism while swimming submerged by combining elements of both dive and exercise responses, with one or the other predominating depending on the level of exertion.     

Regional heterothermy and conservation of core temperature in emperor penguins diving under sea ice

Temperatures were recorded at several body sites in emperor penguins (Aptenodytes forsteri) diving at an isolated dive hole in order to document temperature profiles during diving and to evaluate the role of hypothermia in this well-studied model of penguin diving physiology. Grand mean temperatures (+/-S.E.) in central body sites during dives were: stomach: 37.1+/-0.2 degrees C (n=101 dives in five birds), pectoral muscle: 37.8+/-0.1 degrees C (n=71 dives in three birds) and axillary/brachial veins: 37.9+/-0.1 degrees C (n=97 dives in three birds). Mean diving temperature and duration correlated negatively at only one site in one bird (femoral vein, r=-0.59, P<0.05; range <1 degrees C). In contrast, grand mean temperatures in the wing vein, foot vein and lumbar subcutaneous tissue during dives were 7.6+/-0.7 degrees C (n=157 dives in three birds), 20.2+/-1.2 degrees C (n=69 in three birds) and 35.2+/-0.2 degrees C (n=261 in six birds), respectively. Mean limb temperature during dives negatively correlated with diving duration in all six birds (r=-0.29 to -0.60, P<0.05). In two of six birds, mean diving subcutaneous temperature negatively correlated with diving duration (r=-0.49 and -0.78, P<0.05). Sub-feather temperatures decreased from 31 to 35 degrees C during rest periods to a grand mean of 15.0+/-0.7 degrees C during 68 dives of three birds; mean diving temperature and duration correlated negatively in one bird (r=-0.42, P<0.05). In general, pectoral, deep venous and even stomach temperatures during diving reflected previously measured vena caval temperatures of 37-39 degrees C more closely than the anterior abdominal temperatures (19-30 degrees C) recently recorded in diving emperors. Although prey ingestion can result in cooling in the stomach, these findings and the lack of negative correlations between internal temperatures and diving duration do not support a role for hypothermia-induced metabolic suppression of the abdominal organs as a mechanism of extension of aerobic dive time in emperor penguins diving at the isolated dive hole. Such high temperatures within the body and the observed decreases in limb, anterior abdomen, subcutaneous and sub-feather temperatures are consistent with preservation of core temperature and cooling of an outer body shell secondary to peripheral vasoconstriction, decreased insulation of the feather layer, and conductive/convective heat loss to the water environment during the diving of these emperor penguins.     

A review of the multi-level adaptations for maximizing aerobic dive duration in marine mammals: from biochemistry to behavior

Marine mammals exhibit multi-level adaptations, from cellular biochemistry to behavior, that maximize aerobic dive duration. A dive response during aerobic dives enables the efficient use of blood and muscle oxygen stores, but it is exercise modulated to maximize the aerobic dive limit at different levels of exertion. Blood volume and concentrations of blood hemoglobin and muscle myoglobin are elevated and serve as a significant oxygen store that increases aerobic dive duration. However, myoglobin is not homogeneously distributed in the locomotory muscles and is highest in areas that produce greater force and consume more oxygen during aerobic swimming. Muscle fibers are primarily fast and slow twitch oxidative with elevated mitochondrial volume densities and enhanced oxidative enzyme activities that are highest in areas that produce more force generation. Most of the muscle mitochondria are interfibriller and homogeneously distributed. This reduces the diffusion distance between mitochondria and helps maintain aerobic metabolism under hypoxic conditions. Mitochondrial volume densities and oxidative enzyme activities are also elevated in certain organs such as liver, kidneys, and stomach. Hepatic and renal function along with digestion and assimilation continue during aerobic dives to maintain physiological homeostasis. Most ATP production comes from aerobic fat metabolism in carnivorous marine mammals. Glucose is derived mostly from gluconeogenesis and is conserved for tissues such as red blood cells and the central nervous system. Marine mammals minimize the energetic cost of swimming and diving through body streamlining, efficient, lift-based propulsive appendages, and cost-efficient modes of locomotion that reduce drag and take advantage of changes in buoyancy with depth. Most dives are within the animal’s aerobic dive limit, which maximizes time underwater and minimizes recovery time at the surface. The result of these adaptations is increased breath-hold duration and enhanced foraging ability that maximizes energy intake and minimizes energy output while making aerobic dives to depth. These adaptations are the long, evolutionary legacy of an aquatic lifestyle that directly affects the fitness of marine mammal species for different diving abilities and environments.     

Heart rates and gas exchange in the Amazonian Manatee (Trichechus inunguis) in relation to diving

Unrestrained Amazonian manatees (Trichechus inunguis) maintained a constant heart rate during diving and exhibited a slight tachycardia during breathing. 'Forcing' the manatees to dive caused a marked bradycardia. They exhibited a more pronounced tachycardia during breathing after 'forced' dives and hyperventilated during recovery dives. Manatees are capable of dives exceeding 10 min duration without having to resport to anaerobic metabolism, and even after 10 min dives recover within 3-4 short dives. The ability of manatees to make long dives, in spite of relatively poor O2 stores, is due to their low metabolic rate, while the rapid recovery is aided by their high CO2 stores which minimizes CO2 storage in the body. In manatees the changes in alveolar O2 and CO2 pressure (PAO2 and PACO2) in relation to dive time are slower and more variable than in other marine mammals. The lower rate of change is probably due to the manatees' reduced metabolic rate, while the greater variability is due to their breathing pattern, in which both ventilation and body gas stores influence alveolar gases.     

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.     

Assessing the validity of the accelerometry technique for estimating the energy expenditure of diving double-crested cormorants Phalacrocorax auritus

Over the past few years, acceleration-data loggers have been used to provide calibrated proxies of energy expenditure: the accelerometry technique. Relationships between rate of oxygen consumption and a derivation of acceleration data termed "overall dynamic body acceleration" (ODBA) have now been generated for a range of species, including birds, mammals, and amphibians. In this study, we examine the utility of the accelerometry technique for estimating the energy expended by double-crested cormorants Phalacrocorax auritus to undertake a dive cycle (i.e., a dive and the subsequent pause at the surface before another dive). The results show that ODBA does not calibrate with energy expenditure in diving cormorants, where energy expenditure is calculated from measures of oxygen uptake during surface periods between dives. The possible explanations include reasons why energy expenditure may not relate to ODBA but also reasons why oxygen uptake between dives may not accurately represent energy expenditure during a dive cycle.     

Diving behaviour and diet of the blue-eyed shag at South Georgia

Summary This paper describes a concurrent investigation of individual variation in diet, diving patterns and performance of blue-eyed shags Phalacrocorax atriceps breeding at South Georgia. Within one day individual shags exhibited one of three foraging strategies: short diving (4 birds, all dives 120 s) and mixed diving (15 birds, predominantly long but with a few short dives). The mean number of dives per day was significantly higher in shags that only made short dives (mean=172.0, SE=43.2) than birds with a mixed diving strategy (mean=40.5, SE=4.7) and birds that made only long dives (mean=30.8, SE=1.8). Diet was assessed using hard remains recovered from pellets regurgitated by the shags. Small nototheniid fish (c. 10 kJ per item) were by far the commonest prey but most pellets contained additional items. The frequency of pellets with additional items of higher energy value than nototheniid fish (10.c. 900 kJ per item), lower energy value (>1–10 kJ per item) and both higher and lower energy items was strikingly similar to the frequency of shags making long, short and both long and short dives respectively. Predicted aerobic dive limits suggested that during long dives, blue-eyed shags were probably sustained by anaerobic metabolism. Models of prey capture rates demonstrated that for both long and short diving, many items must be caught per dive when birds are feeding on prey at the lower end of the energy range. Predicted capture rates for the commonest recorded prey (small fish) differ markedly between the two diving strategies.     

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.     

Anaerobic metabolism in human skeletal muscle during short-term, intense activity.

The ability of human skeletal muscle to provide anaerobically derived ATP during short-term, intense activity is examined. The paper emphasizes the information obtained from direct measurements of substrates, intermediates, and products of the pathways in muscle that provide anaerobically derived ATP. The capacity of muscle to provide ATP via anaerobic pathways is approximately 370 mmol/kg dry muscle (dm) during dynamic exercise lasting approximately 3 min. Anaerobic glycolysis provided approximately 80%, phosphocreatine (PCr) degradation approximately 16%, and depletion of the ATP store approximately 4% of the total ATP provided. When the blood flow to the working muscles is reduced or occluded, the anaerobic capacity decreases to approximately 300 mmol/kg dm. This reduction is due to a lower glycolytic capacity associated with an inability to remove lactate from the muscles. Directly measured maximal rates of anaerobically derived ATP provision from PCr degradation and glycolysis during intense muscular activity are each approximately 9-10 mmol.kg-1 dm.s-1. Evidence suggests that both of these pathways are activated instantaneously at the onset of maximal activity. Spring training does little to the capacity or rates of the pathways, although a 10-20% increase in glycolytic ATP provision has been reported. The only study comparing direct and indirect estimates of the anaerobic capacity in humans suggests that O2 deficit measured at the mouth accurately predicts the anaerobic capacity of a single muscle group and that O2 debt does not. There are many unresolved issues regarding the capacity of the PCr and glycogenolytic--glycolytic systems to provide ATP during short-term intense muscular activity in humans.(ABSTRACT TRUNCATED AT 250 WORDS)     

When cormorants go fishing: the differing costs of hunting for sedentary and motile prey

Cormorants hunt both benthic (sedentary) and pelagic (motile) prey but it is not known if the energy costs of foraging on these prey differ. We used respirometry to measure the costs of diving in double-crested cormorants (Phalacrocorax auritus) foraging either for sedentary (fish pieces) or motile (juvenile salmon) prey in a deep dive tank. Short dives for sedentary prey were more expensive than dives of similar duration for motile prey (e.g. 20% higher for a 10s dive) whereas the reverse was true for long dives (i.e. long dives for motile prey were more expensive than for sedentary prey). Across dives of all durations, the foraging phase of the dive was more expensive when the birds hunted motile prey, presumably due to pursuit costs. The period of descent in all the dives undertaken appears to have been more expensive when the birds foraged on sedentary prey, probably due to a higher swimming speed during this period.     

The optimal allocation of time and respiratory metabolism over the dive cycle

Because anaerobic metabolism is much less efficient than aerobicmetabolism in supplying energy, it is widely believed that diversrely predominantly on aerobic metabolism for diving. In thispaper, a time budget model, which assumes that the diver canuse either completely aerobic or partially aerobic metabolismwith additional anaerobic metabolism for diving, is developedand is used to make predictions about patterns in optimal allocationof time and respiratory metabolism during the dive cycle. Theresults derived from the model are (1) a diver that can varythe ratio of energy supplied anaerobically to total energy spentduring dive time is favored by natural selection, but the patternsof time allocation over the dive cycle by the diver do not differ fromthose of a diver that cannot vary the ratio. (2) Even if itis assumed that divers switch their metabolism for diving, anobvious upturn in the surface time with respect to dive timedoes not occur at the aerobic dive limit (ADL) but occurs beyondthe ADL. (3) Use of additional anaerobic metabolism can be favoredfor dives shorter than the ADL. These findings provide a usefulguide to understanding the factors that limit diving behavior.     

Diving in shallow water: the foraging ecology of darters (Aves: Anhingidae)

Diving birds have to overcome buoyancy, especially when diving in shallow water. Darters and anhingas (Anhingidae) are specialist shallow-water divers, with adaptations for reducing their buoyancy. Compared to closely-related cormorants (Phalacrocoracidae), darters have fully wettable plumage, smaller air sacs and denser bones. A previous study of darter diving behaviour reported no relationship between dive duration and water depth, contrary to optimal dive models. In this study I provide more extensive observations of African darters Anhinga melanogaster rufa diving in water<5 m deep at two sites. Dive duration increases with water depth at both sites, but the relationship is weak. Dives were longer than dives by cormorants in water of similar depth (max 108 s in water 2.5 m deep), with dives of up to 68 s observed in water<0.5 m deep. Initial dives in a bout were shorter than expected, possibly because their plumage was not fully saturated. Dive efficiency (dive:rest ratio) was 5–6, greater than cormorants (2.7±0.4 for 18 species) and other families of diving birds (average 0.2–4.3). Post-dive recovery periods increased with dive duration, but only slowly, resulting in a strong increase in efficiency with dive duration. All dives are likely to fall within the theoretical anaerobic dive limit. Foraging bouts were short (17.8±4.3 min) compared to cormorants, with birds spending 80±5% of time underwater. Darters take advantage of their low buoyancy to forage efficiently in shallow water, and their slow, stealthy dives are qualitatively different from those of other diving birds. However, they are forced to limit the duration of foraging bouts by increased thermoregulatory costs associated with wettable plumage.     

Cardiac output and muscle blood flow during rest-associated apneas of elephant seals

In order to evaluate hemodynamics and blood flow during rest-associated apnea in young elephant seals (Mirounga angustirostris), cardiac outputs (CO, thermodilution), heart rates (HR), and muscle blood flow (MBF, laser Doppler flowmetry) were measured. Mean apneic COs and HRs of three seals were 46% and 39% less than eupneic values, respectively (2.1+/-0.3 vs. 4.0+/-0.1 mL kg(-1) s(-1), and 54+/-6 vs. 89+/-14 beats min(-1)). The mean apneic stroke volume (SV) was not significantly different from the eupneic value (2.3+/-0.2 vs. 2.7+/-0.5 mL kg(-1)). Mean apneic MBF of three seals was 51% of the eupneic value. The decline in MBF during apnea was gradual, and variable in both rate and magnitude. In contrast to values previously documented in seals during forced submersions (FS), CO and SV during rest-associated apneas were maintained at levels characteristic of previously published values in similarly-sized terrestrial mammals at rest. Apneic COs of such magnitude and incomplete muscle ischemia during the apnea suggest that (1) most organs are not ischemic during rest-associated apneas, (2) the blood O(2) depletion rate is greater during rest-associated apneas than during FS, and (3) the blood O(2) store is not completely isolated from muscle during rest-associated apneas.     

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.     

A phylogenetic analysis of the allometry of diving

The oxygen store/usage hypothesis suggests that larger animals are able to dive for longer and hence deeper because oxygen storage scales isometrically with body mass, whereas oxygen usage scales allometrically with an exponent <1 (typically 0.67-0.75). Previous tests of the allometry of diving tend to reject this hypothesis, but they are based on restricted data sets or invalid statistical analyses (which assume that every species provides independent information). Here we apply information-theoretic statistical methods that are phylogenetically informed to a large data set on diving variables for birds and mammals to describe the allometry of diving. Body mass is strongly related to all dive variables except dive:pause ratio. We demonstrate that many diving variables covary strongly with body mass and that they have allometric exponents close to 0.33. Thus, our results fail to falsify the oxygen store/usage hypothesis. The allometric relationships for most diving variables are statistically indistinguishable for birds and mammals, but birds tend to dive deeper than mammals of equivalent mass. The allometric relationships for all diving variables except mean dive duration are also statistically indistinguishable for all major taxonomic groups of divers within birds and mammals, with the exception of the procellariiforms, which, strictly speaking, are not true divers.     

Blood Oxygen Depletion Is Independent of Dive Function in a Deep Diving Vertebrate,the Northern Elephant Seal

Although energetics is fundamental to animal ecology, traditional methods of determining metabolic rate are neither direct nor instantaneous. Recently, continuous blood oxygen (O₂) measurements were used to assess energy expenditure in diving elephant seals (Mirounga angustirostris), demonstrating that an exceptional hypoxemic tolerance and exquisite management of blood O₂ stores underlie the extraordinary diving capability of this consummate diver. As the detailed relationship of energy expenditure and dive behavior remains unknown, we integrated behavior, ecology, and physiology to characterize the costs of different types of dives of elephant seals. Elephant seal dive profiles were analyzed and O₂ utilization was classified according to dive type (overall function of dive: transit, foraging, food processing/rest). This is the first account linking behavior at this level with in vivo blood O₂ measurements in an animal freely diving at sea, allowing us to assess patterns of O₂ utilization and energy expenditure between various behaviors and activities in an animal in the wild. In routine dives of elephant seals, the blood O₂ store was significantly depleted to a similar range irrespective of dive function, suggesting that all dive types have equal costs in terms of blood O₂ depletion. Here, we present the first physiological evidence that all dive types have similarly high blood O₂ demands, supporting an energy balance strategy achieved by devoting one major task to a given dive, thereby separating dive functions into distinct dive types. This strategy may optimize O₂ store utilization and recovery, consequently maximizing time underwater and allowing these animals to take full advantage of their underwater resources. This approach may be important to optimizing energy expenditure throughout a dive bout or at-sea foraging trip and is well suited to the lifestyle of an elephant seal, which spends > 90% of its time at sea submerged making diving its most “natural” state.     

Time/depth usage of Adélie penguins: an approach based on dive angles

Data on the swim speed, dive depth and feeding rates of three Adélie penguins (Pygoscelis adeliae) foraging in summer 1998/1999 in Adélie Land, Antarctica were collected using dorsally-mounted loggers, in tandem with oesophageal temperature sensors. Swim speed could be integrated, together with the rate of change of depth, to determine dive and return-to-surface angles. Overall, birds increased rates of change of depth during commuting phases so that dive angles were steeper in dives terminating at greater depths. Angles of descent and ascent during feeding dives were greater than during non-feeding dives. Variation in the descent angle over time of particular dives was generally less than 10°, but the angles of the ascent phases varied more widely. The importance of selecting the optimum descent and ascent angles with respect to prey exploitation, oxygen stores and time gained in the feeding area over the course of a dive by diving at a steeper angle is discussed.     

The diving paradox: new insights into the role of the dive response in air-breathing vertebrates

When aquatic reptiles, birds and mammals submerge, they typically exhibit a dive response in which breathing ceases, heart rate slows, and blood flow to peripheral tissues is reduced. The profound dive response that occurs during forced submergence sequesters blood oxygen for the brain and heart while allowing peripheral tissues to become anaerobic, thus protecting the animal from immediate asphyxiation. However, the decrease in peripheral blood flow is in direct conflict with the exercise response necessary for supporting muscle metabolism during submerged swimming. In free diving animals, a dive response still occurs, but it is less intense than during forced submergence, and whole-body metabolism remains aerobic. If blood oxygen is not sequestered for brain and heart metabolism during normal diving, then what is the purpose of the dive response? Here, we show that its primary role may be to regulate the degree of hypoxia in skeletal muscle so that blood and muscle oxygen stores can be efficiently used. Paradoxically, the muscles of diving vertebrates must become hypoxic to maximize aerobic dive duration. At the same time, morphological and enzymatic adaptations enhance intracellular oxygen diffusion at low partial pressures of oxygen. Optimizing the use of blood and muscle oxygen stores allows aquatic, air-breathing vertebrates to exercise for prolonged periods while holding their breath.     

Dive bout organization in the Chinstrap penguin at seal island, antarctica

Diving behavior of 2 breeding Chinstrap penguins (Pygoscelis antarctica) was studied focusing first and primarily on dive bouts rather than dives themselves. Analysis of dive bout organization revealed (1) though there are differences between solitary dives and dive bouts in dive duration and dive depth, the first dives of dive bouts do not differ from solitary dives in the dive parameters, (2) mean dive duration during bout correlates positively to both mean dive depth during bout and mean surface interval during bout, while number of dives during bout negatively correlates to both cost (consumed energy) and duration of a dive cycle during bout. These findings suggest the following possibilities on foraging behavior of penguins: (1) their decision to repeat diving depends on the result of the first dive at a site, and the first dives of bouts would tend to be searching or evaluating dives though they would be also successful foraging dives, (2) they repeat diving at a foraging patch until foraging efficiency decrease to a threshold of diminishing returns.     

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.