Hemoglobin concentrations and blood gas tensions of free-diving Weddell seals

Arterial blood gas tensions, pH, and hemoglobin concentrations were measured in four free-diving Weddell seals Leptonychotes weddelli. A microprocessor-controlled sampling system enabled us to obtain 24 single and 31 serial aortic blood samples. The arterial O2 tension (PaO2) at rest [78 +/- 13 (SD) Torr] increased with diving compression to a maximum measured value of 232 Torr and then rapidly decreased to 25-35 Torr. The lowest diving PaO2 we measured was 18 Torr just before the seal surfaced from a 27-min dive. A consistent increase of arterial hemoglobin concentrations from 15.1 +/- 1.10 to 22.4 +/- 1.41 g/100 ml (dives less than 17 min) and to 25.4 +/- 0.79 g/100 ml (dives greater than 17 min) occurred during each dive. We suggest that an extension of the sympathetic outflow of the diving reflex possibly caused profound contraction of the Weddell seal's very large spleen (0.89% of body wt at autopsy), although we have no direct evidence. This contraction may have injected large quantities of red blood cells (2/3 of the total) into the seal's central circulation during diving and allowed arterial O2 content to remain constant for the first 15-18 min of long dives. The increase of arterial CO2 tensions during the dive and the compression increase of arterial N2 tensions were also moderated by injecting red blood cells sequestered at ambient pressure. After each dive circulating red blood cells are oxygenated and rapidly sequestered, possibly in the spleen during the first 15 min of recovery.     

DIVING BEHAVIOR AND PERFORMANCE OF HARBOR PORPOISES, PHOCOENA PHOCOENA, IN FUNKA BAY, HOKKAIDO, JAPAN

In order to monitor the diving behavior of free-ranging cetaceans, microdataloggers, with pre-programmed release mechanisms, were attached to the dorsal fins of two female harbor porpoises ( Phocoena phocoena ) in Funka Bay, Hokkaido, Japan, in 1994. The two loggers were successfully recovered and a total of 141 h of diving data (depth and water temperature in 4,671 dives) was obtained. Both porpoises dived almost continuously, rarely exhibiting long-term rest at the surface. Maximum dive depths were 98.6 m and 70.8 m, respectively, with more than 70% of diving time at 20 m or less. Most shallow dives were V-shaped with no bottom time. The V-shaped dives were significantly shallower in dive depth and shorter in dive duration than U-shaped dives. Descent rate was not constant during a dive. The deeper the dive depths, the faster the mean descent and initial descent rates. This suggests that porpoises have anticipated the depth to which they will dive before initiating the dive itself.     

Artifacts arising from sampling interval in dive depth studies of marine endotherms

Most depth recorders used to study the diving behaviour of polar marine endotherms record depth data at specific time intervals. The length of recording interval can have potentially profound implications for the interpretation of the data. We used data acquired on the diving behaviour of king penguins, Aptenodytes patagonicus, to examine the validity of various analyses routinely conducted on depth data. In our experiments, increasing the sampling interval led to an underestimation of the number of dives performed, an overestimation in mean dive duration and substantial changes in the form of the dive profile. Our analysis indicates that depth data should be recorded at a minimum rate corresponding to 10% of the total dive duration and that conventional dive profile categorization may be inappropriate. Alternatives that are less subjective, and based on curve fits of dive depth versus time, are proposed.     

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.     

TWO APPROACHES TO COMPRESSING AND INTERPRETING TIME-DEPTH INFORMATION AS AS COLLECTED BY TIME-DEPTH RECORDERS AND SATELLITE-LINKED DATA RECORDERS

Time-depth recorders sample information about the three- dimensional behavior of diving animals over time and reduce this into just two dimensions, depth and time. Even so, interpretation of the data may still be difficult because of the volume of data and the detail that remains. Comparison of dive "shape" across individuals, geographical locations, or species presents problems because its analysis may involve subjective judgments or arbitrary distinctions. More constraints may be imposed if a telemetry system is used to transmit the data. Here we present two approaches for dive data compression and analysis. The first (applied before storage and transmission) selects the most important time-depth points in a profile where depth Vs. time trajectories change most significantly. The second (used to either preprocess or postprocess dive information) creates a dimensionless, depth, and duration independent index (TAD), which encapsulates the relevant information from dive profiles on where the diver centers its activity with respect to depth during a dive. Its use facilitates comparison across dives performed at different times or places, within or between individuals or species, irrespective of the duration and depth of their dives. Both can be used to reduce the amount of information sent or stored about dive behavior and can facilitate dive analysis.     

Optimizing methods to estimate zooplankton concentration based on generalized patterns of patchiness inside ballast tanks and ballast water discharges

Zooplankton populations are spatially heterogeneous in nature and inside ship ballast tanks. Sampling methods should take heterogeneity into account, particularly when estimating quantitative variables such as abundance or concentration. It is particularly important to generate unbiased estimates of zooplankton concentration in ballast water when assessing compliance with new international ballast water discharge standards. We measured spatial heterogeneity of zooplankton within ballast water using three sampling methodologies. In‐tank pump samples were collected at fixed depths within the vertical part of the ballast tank (side tank). Vertical net‐haul samples were collected from the upper portion of the tank as a depth‐integrated and historically relevant method. In‐line, time‐integrated samples were collected during ballast discharge by an isokinetic sample probe, likely representing the double bottom part of the ballast tank. The bias and precision associated with each sampling method were evaluated in reference to the estimated average abundance of the entire ballast tank, which was modeled from the data collected by all methods. In‐tank pump samples provided robust evidence for vertical stratification of zooplankton concentration in the side tank. A consistent trend was also observed for in‐line discharge samples, with zooplankton concentration decreasing through time as the ballast tank is being discharged. Sample representativeness, as compared to the tank average, varied depending on the depth or tank volume discharged. In‐line discharge samples provided the least biased and most precise estimate of average tank abundance (having lowest mean squared error) when collected during the time frame of 20%–60% of the tank volume being discharged. Results were consistent across five trips despite differences in ballast water source, season, and age.     

Is mass loss in Brünnich's guillemots Uria lomvia an adaptation for improved flight performance or improved dive performance?

Breeding Brünnich's guillemots Uria lomvia show stepwise mass loss at the time of hatch. This mass loss has usually been explained as an adaptation to reduce the cost of flight during the chick‐rearing period because flight time increases during that period. It is possible, however, that mass loss also increases dive performance during the chick‐rearing period because time spent diving also increases during that period. Reduced mass could reduce basal metabolic rate or costs associated with buoyancy and therefore increase aerobic dive limit. To examine the role of mass loss in dive behavior, we attached time‐depth‐temperature recorders for 24–48 h to chick‐rearing and incubating Brünnich's guillemots at Coats Island, Nunavut (2005: n=45, 2006: n=40), and recorded body mass before and after each deployment. There was no relationship between mass and dive duration during either incubation or chick‐rearing. Seventeen of the birds we sampled during incubation were resampled during chick‐rearing. For this group, dive duration increased with mass loss between incubation and chick‐rearing (r²=0.67–0.75). Mass loss occurred through reductions in metabolically‐active tissues (liver, bladder) and buoyant tissues (lipids) although muscle and gut mass did not change. Despite the large change in lipids, buoyancy only changed by 0.1%, and mass loss therefore did not have much effect on costs associated with buoyancy. Nonetheless, surface pause duration for a given dive depth decreased during chick‐rearing, supporting the idea that reduced mass led to increased aerobic dive limit through reduced metabolic rate and inertial costs; oxygen stores did not increase. We also attached neutrally (n=9) and negatively (n=11) buoyant handicaps to the legs of adults to assess the effect of artificial mass increases on time budgets. Artificially increasing mass decreased total time spent diving but did not change time spent flying. There was no change in shift length between incubation and chick‐rearing, and therefore no support for the idea that mass loss reflected a change in fasting endurance requirements. An energetic model suggested that the observed mass reduction reduced dive costs by 5–8% and flight costs by 3%. We concluded that mass loss may be as important for increasing dive performance as increasing flight performance.     

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.     

Nitrogen tensions in brachial vein blood of Korean ama divers.

Intravascular bubble formation and symptoms of decompression sickness have been reported during repetitive deep breath-hold diving. Therefore we examined the pattern of blood N2 kinetics during and after repetitive breath-hold diving. To study muscle N2 uptake and release, we measured brachial venous N2 partial pressure (PN2) in nine professional Korean breath-hold divers (ama) during a 3-h diving shift at approximately 4 m seawater depth and up to 4 h after diving. PN2 was determined with the manometric Van Slyke method. Diving time and depth were recorded using a backpack computer-assisted dive longer that allowed calculating the surface-to-depth time ratio to derive the effective depth. With the assumption that forearm muscle N2 kinetics follow the general Haldanian principles of compression and decompression, i.e., forearm muscle is a single compartment with a uniform tissue PN2 equal to venous PN2, PN2 data were fitted to monoexponential functions of time. In the early phase of the diving shift, PN2 rapidly increased to 640 Torr (half time = 6 min) and then slowly declined to baseline levels (half time = 36 min) after the work shift. Peak PN2 levels approximated the alveolar PN2 derived from the effective depth. We conclude that forearm muscle N2 kinetics are well described by a Haldanian single-compartment model. Decompression sickness is theoretically possible in the ama; it did not occur because the absolute PN2 remained low due to the shallow working depth of the ama we studied.     

The diving behaviour of the Manx Shearwater Puffinus puffinus

The diving capabilities of the Procellariformes remain the least understood component of avian diving physiology. Due to their relatively small size, shearwaters may have high oxygen consumption rates during diving relative to their available oxygen stores. Dive performance in this group should be strongly limited by the trade‐off between oxygen consumption and oxygen stores, and shearwaters could be a good model group for testing predictions of dive theory. Many earlier measurements of shearwater dive behaviour relied on observations from the surface or potentially biased technology, and it is only recently that diving behaviour has been observed using electronic recorders for many of the clades within the family. The diving behaviour of Manx Shearwaters Puffinus puffinus breeding in Wales, UK, was studied on a large sample of birds using time–depth–temperature recorders deployed on chick‐rearing shearwaters in July and August over 3 years (2009–2011). Light availability apparently limited diving as dives only occurred between 04:00 and 19:00 h GMT. All individuals routinely dived deeper than traditionally assumed, to a mean maximum depth of 31 m and occasionally down to nearly 55 m. We compiled all available data for a comparison of the dive depth across shearwater species. There was a positive allometric relationship between maximum dive depth and body mass across Puffinus and Ardenna shearwater species, as expected, but only if samples of fewer than two individuals were excluded. The large intra‐specific range in maximum dive depth in our study illustrates that apparent diversity in diving performance across species must be interpreted cautiously.     

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.     

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.     

Diel differences in avoidance of three vertical profile sampling gears by herring larvae

Three plankton sampling systems [pump, Longhurst-Hardy PlanktonRecorder (LHPR) and MOCNESS] were used to obtain vertical profilesof herring larvae off the north-east coast of England in October1986 during the International Herring Larval Survey (IHLS).A factorial sampling design allowed a comparison of the numberof herring larvae caught by depth (four levels), time of day(three levels) and gear (three levels). The size compositionof herring larvae taken in the different gears has also beenexamined. Catches taken at night were greater than those takenduring the day throughout the water column, particularly withlarger larvae. This indicates that visual avoidance of all gearsis an important factor, which increases as the larvae grow bigger.LHPR night catches were greater than in the MOCNESS and fargreater than in the pump, which indicates that avoidance isalso due to senses other than vision. Examination of the 1981–1986International Herring Larvae Survey catch data showed that night-timecatches were similar to daytime catches for herring larvae <10mm long, but night time catches caught progressively more asthe length of the larvae increased to 24 mm.     

Optimal diving under the risk of predation

Many air-breathing aquatic foragers may be killed by aerial or subsurface predators while recovering oxygen at the surface; yet the influence of predation risk on time allocation during dive cycles is little known in spite of numerous studies on optimal diving. We modeled diving behavior under the risk of predation at the surface. The relationship between time spent at the surface and the risk of death is predicted to influence the optimal surface interval, regardless of whether foragers accumulate energy at a constant rate while at the food patch, deplete food resources over the course of the dive, or must search for food during the dive. When instantaneous predation risk during a single surface interval decreases with time spent at the surface, a diver should increase its surface interval relative to that which maximizes energy intake, thereby increasing dive durations and reducing the number of surfacings per foraging bout. When instantaneous risk over a single surface interval does not change or increases with increasing time at the surface, divers should decrease their surface interval (and consequently their dive duration) relative to that which maximizes energy intake resulting in more dives per foraging bout. The fitness consequences of selecting a suboptimal surface interval vary with the risk function and the way divers harvest energy when at depth. Finally, predation risk during surface intervals should have important consequences for habitat selection and other aspects of the behavioral ecology of air-breathing aquatic organisms.     

Swim speed of free-ranging Adélie penguins Pygoscelis adeliae and its relation to the maximum depth of dives

Swim speed and depth utilization were recorded at a sampling rate of 1 Hz in 14 free-ranging Adélie penguins in Adélie Land, Antarctica during the austral summers of 1996/1997 and 1998/1999. The average swim speeds during the descent, bottom and ascent phases of dives were independent of the maximum depth, while the variability in swim speed decreased with increasing maximum depth, reflecting the physiological constraints of diving. Descent speed, which varied less with maximum depth than speeds measured during other parts of dives, was significantly different among birds. In addition to the speed analysis, a new category of dive profiles with a flat bottom phase and an extremely reduced swim speed is reported. The probable benthic nature of such dives is discussed.     

Pattern and depth of dives in Northern elephant seals, Mirounga angustirostris

A time-depth recorder was attached to a female Northern elephant seal at the end of her lactation fast before she entered the sea to feed. The animal dived continuously during its first 11 days at sea, the period recorded for a total of 653 dives. Mean dive time was 21 min, with the longest submersion lasting 32 min. Mean surface interval between dives was 3 min, resulting in a total surface time of 11 %. Mean dive depth was 333 m and the deepest dive was 630 m, the deepest ever recorded for a pinniped. A epth histogram recorder attached to another female yielded a similar frequency distribution of dive depths.     

Diving heart rate development in postnatal harbour seals, Phoca vitulina

Harbour seals, Phoca vitulina, dive from birth, providing a means of mapping the development of the diving response, and so our objective was to investigate the postpartum development of diving bradycardia. The study was conducted May-July 2000 and 2001 in the St. Lawrence River Estuary (48 degrees 41'N, 68 degrees 01'W). Both depth and heart rate (HR) were remotely recorded during 86,931 dives (ages 2-42 d, n = 15) and only depth for an additional 20,300 dives (combined data covered newborn to 60 d, n = 20). The mean dive depth and mean dive durations were conservative during nursing (2.1 +/- 0.1 m and 0.57 +/- 0.01 min, range = 0-30.9 m and 0-5.9 min, respectively). The HR of neonatal pups during submersion was bimodal, but as days passed, the milder of the two diving HRs disappeared from their diving HR record. By 15 d of age, most of the dive time was spent at the lower diving bradycardia rate. Additionally, this study shows that pups are born with the ability to maintain the lower, more fully developed dive bradycardia during focused diving but do not do so during shorter routine dives.     

Dive types and circadian behaviour patterns of Saimaa ringed seals<Emphasis Type="Italic">Phoca hispida saimensis</Emphasis> during the open-water season

Diving and circadian behaviour patterns of 7 free-ranging Saimaa ringed sealsPhoca hispida saimensis Nordquist, 1899 were examined by VHF-radiotelemetry during open-water seasons between May and November in Lake Saimaa, eastern Finland. The mean recorded dive duration ranged from 2.8 to 6.5 min, with a maximum of 21 min. The mean dive depth ranged from 9.8 to 15.7 m, with maximum of 39.6 m. The maximum dive depth of each seal was limited by water depth in the study area. The dive depths were positively correlated with dive duration and body mass of the seal. Five different dive types were defined, as based on their depth-time characteristics, each falling into one of the three functional categories: travelling, feeding, and resting. Long duration diving bouts occurred mostly at night and were presumed to be resting dives. Saimaa ringed seals exhibited a circadian pattern of haul-out behaviour that shifted seasonally. During molting (May–June) the seals hauled-out both day and night, but later in summer haul-out was more frequent at night.     

DIVING BEHAVIOR OF SUBADULT AND ADULT HARBOR SEALS IN PRINCE WILLIAM SOUND, ALASKA

Satellite-linked depth recorders (SDRs) were attached to 47 harbor seals in Prince William Sound, Alaska, during 1992–1996. Parameters describing diving effort, diving focus, and focal depth (depth bin to which diving was focused) were calculated from binned data on maximum dive depth and time spent at depth, and analyzed using repeated-measures mixed models. This analysis method accounted for individual variability, temporal autocorrelation, and the binned nature of SDR data, which are often ignored using standard statistical techniques. Results indicated that diving effort remained steady from September to April, when seals spent 68%-75% of their overall time in the water. Time spent in the water declined to 60% in May and to about 40% in July. Seals spent the most time in the water at night and the least in the morning. The diving of all seals in all months was highly focused. Overall, diving was focused to one depth bin approximately 75% of the time. Diving was more focused for females than for males and subadults. Focal dive depth was deepest in winter and shallowest during May-July. Focal depth and diving focus varied by region. Collinearity between month and region in the focal depth model suggests that seals move in winter to regions where prey are found deeper in the water column. Variations in diving behavior presumably result from combinations of regional bathymetry, seasonal cycles in type or depth distribution of prey, and seal life-cycle events such as reproduction and molting.