pH regulation of calcium efflux by turtle liver mitochondria

1. Turtle liver mitochondria are capable of taking up calcium in a pattern similar to that described from rat liver mitochondria. 2. Turtle liver mitochondria also possess a system for calcium efflux which is extremely sensitive to changes in extramitochondrial pH. A decrease in extramitochondrial pH by addition of HCl or by gassing with CO2 caused a rapid release of calcium. 3. The profound changes in pH and pCO2 during deep diving likely affects Ca efflux from mitochondria in the turtle liver.     

Latitudinal range influences the seasonal variation in the foraging behavior of marine top predators

Non-migratory resident species should be capable of modifying their foraging behavior to accommodate changes in prey abundance and availability associated with a changing environment. Populations that are better adapted to change will have higher foraging success and greater potential for survival in the face of climate change. We studied two species of resident central place foragers from temperate and equatorial regions with differing population trends and prey availability associated to season, the California sea lion (Zalophus californianus) (CSL) whose population is increasing and the endangered Galapagos sea lion (Zalophus wollebaeki) (GSL) whose population is declining. To determine their response to environmental change, we studied and compared their diving behavior using time-depth recorders and satellite location tags and their diet by measuring C and N isotope ratios during a warm and a cold season. Based on latitudinal differences in oceanographic productivity, we hypothesized that the seasonal variation in foraging behavior would differ for these two species. CSL exhibited greater seasonal variability in their foraging behavior as seen in changes to their diving behavior, foraging areas and diet between seasons. Conversely, GSL did not change their diving behavior between seasons, presenting three foraging strategies (shallow, deep and bottom divers) during both. GSL exhibited greater dive and foraging effort than CSL. We suggest that during the warm and less productive season a greater range of foraging behaviors in CSL was associated with greater competition for prey, which relaxed during the cold season when resource availability was greater. GSL foraging specialization suggests that resources are limited throughout the year due to lower primary production and lower seasonal variation in productivity compared to CSL. These latitudinal differences influence their foraging success, pup survival and population growth reflected in contrasting population trends in which CSL are more successful and potentially more resilient to climate change.     

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.     

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.     

First records of sea snakes (Elapidae: Hydrophiinae) diving to the mesopelagic zone (>200 m)

Viviparous sea snakes (Elapidae: Hydrophiinae) are fully marine reptiles distributed in the tropical and subtropical waters of the Indian and Pacific Oceans. Their known maximum diving depth ranges between 50 and 100 m and this is thought to limit their ecological ranges to shallow habitats. We report two observations, from industry‐owned remotely operated vehicles, of hydrophiine sea snakes swimming and foraging at depths of approximately 250 m in the Browse Basin on Australia's North West Shelf, in 2014 and 2017. These observations show that sea snakes are capable of diving to the dim‐lit, cold‐water mesopelagic zone, also known as the ‘twilight’ zone. These record‐setting dives raise new questions about the thermal tolerances, diving behaviour and ecological requirements of sea snakes. In addition to significantly extending previous diving records for sea snakes, these observations highlight the importance of university‐industry collaboration in surveying understudied deep‐sea habitats.     

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.     

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.     

Foraging ecology of Gentoo Penguins Pygoscelis papua at Macquarie Island during the period of chick care

The diet, diving behaviour, swimming velocity and foraging range of Gentoo Penguins Pygoscelis papua were studied at Macquarie Island during the breeding season in the 1993–1994 austral summer. Gentoo Penguins are considered to be inshore feeders, and at Macquarie Island the diet and estimated foraging ranges supported this. The diet consisted of 91.6% fish and 8.3% squid, by mass. The dominant prey taxa were the fish Gymnoscopelus sp. and Paranotothenia magellanica. A mixture of pelagic and benthic prey was consumed, with a greater proportion of benthic species occurring later in the season. The penguins exhibited a strong diurnal pattern in their diving behaviour. Deep diving (≥30 m) began near sunrise (03.00 h) and finished close to sunset (21.00 h). Diving at night was less common and very shallow (<10 m). Early in the breeding season, dive profiles indicated that birds were probably following vertically migrating pelagic prey through the water column and were foraging in waters over 100 m deep. Later in the season, more uniform, shallower depths were used, suggesting an increase in benthic foraging activity. These changes in dive pattern and depth were consistent with the habitat preferences of prey species found in the diet. Gentoo Penguins swam at 1.04 m per s and had a maximum potential foraging range of about 26 km for single-day trips. They tended to forage within 14 km of the colony, with a mean range of 5.4 km. This range encompassed the deep ocean habitat to the west and east of the island and a shallow area to the north.     

Changes in body temperature in king penguins at sea: the result of fine adjustments in peripheral heat loss?

To investigate thermoregulatory adjustments at sea, body temperatures (the pectoral muscle and the brood patch) and diving behavior were monitored during a foraging trip of several days at sea in six breeding king penguins Aptenodytes patagonicus. During inactive phases at sea (water temperature: 4-7 degrees C), all tissues measured were maintained at normothermic temperatures. The brood patch temperature was maintained at the same values as those measured when brooding on shore (38 degrees C). This high temperature difference causes a significant loss of heat. We hypothesize that high-energy expenditure associated with elevated peripheral temperature when resting at sea is the thermoregulatory cost that a postabsorptive penguin has to face for the restoration of its subcutaneous body fat. During diving, mean pectoral temperature was 37.6 +/- 1.6 degrees C. While being almost normothermic on average, the temperature of the pectoral muscle was still significantly lower than during inactivity in five out of the six birds and underwent temperature drops of up to 5.5 degrees C. Mean brood patch temperature was 29.6 +/- 2.5 degrees C during diving, and temperature decreases of up to 21.6 degrees C were recorded. Interestingly, we observed episodes of brood patch warming during the descent to depth, suggesting that, in some cases, king penguins may perform active thermolysis using the brood patch. It is hypothesized that functional pectoral temperature may be regulated through peripheral adjustments in blood perfusion. These two paradoxical features, i.e., lower temperature of deep tissues during activity and normothermic peripheral tissues while inactive, may highlight the key to the energetics of this diving endotherm while foraging at sea.     

Deep-diving behaviour of the northern bottlenose whale,Hyperoodon ampullatus (Cetacea: Ziphiidae)

Using suction-cup attached time–depth recorder/VHF radio tags, we have obtained the first diving data on northern bottlenose whales (Hyperoodon ampullatus), the first such data on any species within the family Ziphiidae. Two deployments in 1997 on northern bottlenose whales in a submarine canyon off Nova Scotia demonstrated their exceptional diving ability, with dives approximately every 80 min to over 800 m (maximum 1453 m), and up to 70 min in duration. Sonar traces of non-tagged, diving bottlenose whales in 1996 and 1997 suggest that such deep dives are not unusual. This combined evidence leads us to hypothesize that these whales may make greater use of deep portions of the water column than any other mammal so far studied. Many of the recorded dives of the tagged animals were to, or close to, the sea floor, consistent with benthic or bathypelagic foraging. A lack of correlation between dive times and surface intervals suggests that the dives were predominately aerobic.     

Individual Foraging Strategies Reveal Niche Overlap between Endangered Galapagos Pinnipeds

Most competition studies between species are conducted from a population-level approach. Few studies have examined inter-specific competition in conjunction with intra-specific competition, with an individual-based approach. To our knowledge, none has been conducted on marine top predators. Sympatric Galapagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki) share similar geographic habitats and potentially compete. We studied their foraging niche overlap at Cabo Douglas, Fernandina Island from simultaneously collected dive and movement data to examine spatial and temporal inter- and intra-specific competition. Sea lions exhibited 3 foraging strategies (shallow, intermediate and deep) indicating intra-specific competition. Fur seals exhibited one foraging strategy, diving predominantly at night, between 0–80 m depth and mostly at 19–22 h. Most sea lion dives also occurred at night (63%), between 0–40 m, within fur seals'' diving depth range. 34% of sea lions night dives occurred at 19–22 h, when fur seals dived the most, but most of them occurred at dawn and dusk, when fur seals exhibited the least amount of dives. Fur seals and sea lions foraging behavior overlapped at 19 and 21 h between 0–30 m depths. Sea lions from the deep diving strategy exhibited the greatest foraging overlap with fur seals, in time (19 h), depth during overlapping time (21–24 m), and foraging range (37.7%). Fur seals foraging range was larger. Cabo Douglas northwest coastal area, region of highest diving density, is a foraging “hot spot” for both species. Fur seals and sea lions foraging niche overlap occurred, but segregation also occurred; fur seals primarily dived at night, while sea lions exhibited night and day diving. Both species exploited depths and areas exclusive to their species. Niche breadth generally increases with environmental uncertainty and decreased productivity. Potential competition between these species could be greater during warmer periods when prey availability is reduced.     

Foraging by deep-diving birds is not constrained by an aerobic diving limit: a model of avian depth-dependent diving metabolic rate

The theoretical aerobic diving limit (tADL) specifies the duration of a dive after which oxygen reserves available for diving are depleted. The tADL has been calculated by dividing the available oxygen stores by the diving metabolic rate (DMR). Contrary to diving mammals, most diving birds examined to date exceed the tADL by a large margin. This discrepancy between observation and theory has engendered two alternative explanations suggesting that dive duration is extended either anaerobically or by depressing aerobic metabolism. Current formulations of tADL uncritically assume that DMR is independent of depth. However, diving birds differ from other vertebrate divers by having a larger respiratory system volume and by retaining air in their plumage while diving, thereby elevating buoyancy. Because air compresses with depth, diving power requirement decreases with depth. Following this principle, we modeled DMR to depth for Adelie and little penguins and reformulated the tADL accordingly. The model's results suggest that < approximately 5% of natural dives by Adelie penguins exceed the reformulated tADL(d), or maximal aerobic depth, and none in the more buoyant little penguin. These data suggest that, for both small and large species, deep diving birds rarely if ever exceed tADL(d).     

Diving activity of migrating silver eel with and without Anguillicola crassus infection

Infection with the swim bladder nematode Anguillicola crassus has been hypothesised to threaten the spawning migration success of the endangered European eel (Anguilla anguilla). To examine this assumption, we compared the swimming behaviour of one Anguillicola crassus infested eel in the North Sea and three parasite‐free eels in the Baltic using data recovered from data storage tags attached to migrating silver eels. In both areas, eel activity was characterized by frequent diving behaviour throughout the water column during the night, with reduced activity during the day. Despite substantial damage of the swim bladder, the behaviour of the infested eel from the North Sea was within the same range of migrating and diving activity parameters as the three parasite‐free eels from the Baltic Sea. All eels had a similar frequency distribution of descent or ascent speeds and a similar average horizontal migration speed. The diving speeds and dive ranges exclude the possibility that the eels were in continuous hydrostatic equilibrium during their migrations and suggests therefore that the role of the swim bladder in vertical migration is likely to be more complex than currently thought. Our results suggest that eels infested by Anguillicola crassus are capable of diving in a similar manner to uninfested eels during the first stretch of their spawning migration.     

Deep-diving of Atlantic salmon (<Emphasis Type="Italic">Salmo salar</Emphasis>) during their marine feeding migrations

Data from seven data storage tags recovered from Atlantic salmon marked as smolts were analyzed for depth movements and patterns of deep diving during the marine migration. The salmon mostly stayed at the surface and showed diurnal activity especially from autumn until spring. During the first months at sea the salmon stayed at shallower depths (<100 m). The salmon took short deep dives (>100 m), that were rare or absent during the first summer at sea but increased in frequency and duration especially in late winter. The maximum depth of the dives varied from 419 to 1187 m. Most of dives were short, (<5 h) but could last up to 33 h. The duration of dives increased in late winter until spring and the overall depth and maximum depth per dive increased exponentially over time. The initiation of the dives was more common in evenings and at night, suggesting nocturnal diving. We hypothesized that deep diving is related to feeding of salmon as mesopelagic fish can be important food for salmon during winter.     

Effects of age and body mass on development of diving capabilities of gray seal pups: costs and benefits of the postweaning fast

Development of adequate diving capabilities is crucial for survival of seal pups and may depend on age and body size. We tracked the diving behavior of 20 gray seal pups during their first 3 mo at sea using satellite relay data loggers. We employed quantile analysis to track upper limits of dive duration and percentage time spent diving, and lower limits of surface intervals. When pups first left the breeding colony, extreme (ninety-fifth percentile) dive duration and percentage time spent diving were positively correlated with age, but not mass, at departure. Extreme dive durations and percentage time spent diving peaked at [Formula: see text] d of age at values comparable with those of adults, but were not sustained. Greater peaks in extreme percentage time spent diving occurred in pups that had higher initial values, were older at their peak, and were heavier at departure. Pups that were smaller and less capable divers when they left the colony improved extreme dive durations and percentage time spent diving more rapidly, once they were at sea. Minimum survival time correlated positively with departure mass. Pups that were heavier at weaning thus benefitted from being both larger and older at departure, but smaller pups faced a trade-off. While age at departure had a positive effect on early dive performance, departure mass impacted on peak percentage time spent diving and longer-term survival. We speculate that once small pups have attained a minimum degree of physiological development to support diving, they would benefit by leaving the colony when younger but larger to maximize limited fuel reserves, rather than undergoing further maturation on land away from potential food resources, because poor divers may be able to "catch up" once at sea.     

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.     

Diving behavior in a free‐living,semi‐aquatic herbivore,the Eurasian beaver Castor fiber

Semi‐aquatic mammals have secondarily returned to the aquatic environment, although they spend a major part of their life operating in air. Moving both on land, as well as in, and under water is challenging because such species are considered to be imperfectly adapted to both environments. We deployed accelerometers combined with a depth sensor to study the diving behavior of 12 free‐living Eurasian beavers Castor fiber in southeast Norway between 2009 and 2011 to examine the extent to which beavers conformed with mass‐dependent dive capacities, expecting them to be poorer than wholly aquatic species. Dives were generally shallow (<1 m) and of short duration (<30 s), suggesting that the majority of dives were aerobic. Dive parameters such as maximum diving depth, dive duration, and bottom phase duration were related to the effort during different dive phases and the maximum depth reached. During the descent, mean vectorial dynamic body acceleration (VeDBA—a proxy for movement power) was highest near the surface, probably due to increased upthrust linked to fur‐ and lung‐associated air. Inconsistently though, mean VeDBA underwater was highest during the ascent when this air would be expected to help drive the animals back to the surface. Higher movement costs during ascents may arise from transporting materials up, the air bubbling out of the fur, and/or the animals’ exhaling during the bottom phase of the dive. In a manner similar to other homeotherms, beavers extended both dive and bottom phase durations with diving depth. Deeper dives tended to have a longer bottom phase, although its duration was shortened with increased VeDBA during the bottom phase. Water temperature did not affect diving behavior. Overall, the beavers’ dive profile (depth, duration) was similar to other semi‐aquatic freshwater divers. However, beavers dived for only 2.8% of their active time, presumably because they do not rely on diving for food acquisition.     

Warm eyes provide superior vision in swordfishes

Large and powerful ocean predators such as swordfishes, some tunas, and several shark species are unique among fishes in that they are capable of maintaining elevated body temperatures (endothermy) when hunting for prey in deep and cold water . In these animals, warming the central nervous system and the eyes is the one common feature of this energetically costly adaptation . In the swordfish (Xiphias gladius), a highly specialized heating system located in an extraocular muscle specifically warms the eyes and brain up to 10 degrees C-15 degrees C above ambient water temperatures . Although the function of neural warming in fishes has been the subject of considerable speculation , the biological significance of this unusual ability has until now remained unknown. We show here that warming the retina significantly improves temporal resolution, and hence the detection of rapid motion, in fast-swimming predatory fishes such as the swordfish. Depending on diving depth, temporal resolution can be more than ten times greater in these fishes than in fishes with eyes at the same temperature as the surrounding water. The enhanced temporal resolution allowed by heated eyes provides warm-blooded and highly visual oceanic predators, such as swordfishes, tunas, and sharks, with a crucial advantage over their agile, cold-blooded prey.     

A thorough and quantified method for classifying seabird diving behaviour

Time-depth recorders are commonly deployed on diving animals to obtain information on their aquatic behaviour. The recorded data provide a 2D profile of diving activity. As analyses of diving behaviour from such profiles have become more complex, these analyses have often suffered from a lack of consistency and rigour. There is a growing need for a simple, comparative method to classify diving behaviour thoroughly and quantitatively. Here, a new approach to the classification of the dive profiles of penguins is described, which probably has applicability for many other diving predators as well. This simple approach uses a small, coherent set of criteria to classify behaviours in a detailed and quantified manner, and with relative objectivity. Classification of diving behaviour is possible from the temporal scale of a wiggle within a dive to the scale of a bout of dives. The new method will make comparisons between species easier and clearer because these comparisons will be undertaken within a consistent, more objective framework.     

Differential antioxidant protection in tissues from marine mammals with distinct diving capacities. Shallow/short vs. deep/long divers

The diving response in marine mammals results in bradycardia and peripheral vasoconstriction, with blood flow redistributing preferentially to nervous and cardiac tissues. Therefore, some tissues are rendered ischemic during a dive; with the first breath after a dive, blood flow to all tissues is reestablished. In terrestrial mammals, reactive oxygen species (ROS) production increases in response to ischemia/reperfusion and oxidative damage can occur. The capacity of marine mammals to tolerate repeated ischemia/reperfusion cycles associated with diving appears to be due to an enhanced antioxidant system. However, it is not known if diving depth and/or duration elicit differences in tissue capacity to produce ROS and antioxidant defenses in marine mammals. The objective of this study was to analyze ROS production, antioxidant defenses and oxidative damage in marine mammal species that perform shallow/short vs. deep/long dives. We measured production of superoxide radical (O₂^??), oxidative damage to lipids and proteins, activity of antioxidant enzymes, and glutathione levels in tissues from shallow/short divers (Tursiops truncatus) and deep/long divers (Kogia spp.). We found that differences between the diving capacity of dolphins and Kogia spp. are reflected in O₂^?? production and antioxidant levels. These differences suggest that shallow/short and deep/long divers have distinct mechanisms to successfully maintain redox balance.