The development of an intermediate‐duration tag to characterize the diving behavior of large whales

The development of high‐resolution archival tag technologies has revolutionized our understanding of diving behavior in marine taxa such as sharks, turtles, and seals during their wide‐ranging movements. However, similar applications for large whales have lagged behind due to the difficulty of keeping tags on the animals for extended periods of time. Here, we present a novel configuration of a transdermally attached biologging device called the Advanced Dive Behavior (ADB) tag. The ADB tag contains sensors that record hydrostatic pressure, three‐axis accelerometers, magnetometers, water temperature, and light level, all sampled at 1 Hz. The ADB tag also collects Fastloc GPS locations and can send dive summary data through Service Argos, while staying attached to a whale for typical periods of 3–7 weeks before releasing for recovery and subsequent data download. ADB tags were deployed on sperm whales (Physeter macrocephalus; N = 46), blue whales (Balaenoptera musculus; N = 8), and fin whales (B. physalus; N = 5) from 2007 to 2015, resulting in attachment durations from 0 to 49.6 days, and recording 31 to 2,539 GPS locations and 27 to 2,918 dives per deployment. Archived dive profiles matched well with published dive shapes of each species from short‐term records. For blue and fin whales, feeding lunges were detected using peaks in accelerometer data and matched corresponding vertical excursions in the depth record. In sperm whales, rapid orientation changes in the accelerometer data, often during the bottom phase of dives, were likely related to prey pursuit, representing a relative measure of foraging effort. Sperm whales were documented repeatedly diving to, and likely foraging along, the seafloor. Data from the temperature sensor described the vertical structure of the water column in all three species, extending from the surface to depths >1,600 m. In addition to providing information needed to construct multiweek time budgets, the ADB tag is well suited to studying the effects of anthropogenic sound on whales by allowing for pre‐ and post‐exposure monitoring of the whale's dive behavior. This tag begins to bridge the gap between existing long‐duration but low‐data throughput tags, and short‐duration, high‐resolution data loggers.     

Intraspecific variation in vertical habitat use by tiger sharks (Galeocerdo cuvier) in the western North Atlantic

Tiger sharks (Galeocerdo cuvier) are a wide ranging, potentially keystone predator species that display a variety of horizontal movement patterns, making use of coastal and pelagic waters. Far less, however, is known about their vertical movements and use of the water column. We used pop‐up satellite archival tags with two data sampling rates (high rate and standard rate tags) to investigate the vertical habitat use and diving behavior of tiger sharks tagged on the Puerto Rico–Virgin Islands platform and off Bermuda between 2008 and 2009. Useable data were received from nine of 14 sharks tagged, tracked over a total of 529 days. Sharks spent the majority of their time making yo‐yo dives within the upper 50 m of the water column and considerable time within the upper 5 m of the water column. As a result, sharks typically occupied a narrow daily temperature range (~2°C). Dives to greater than 200 m were common, and all sharks made dives to at least 250 m, with one shark reaching a depth of 828 m. Despite some similarities among individuals, a great deal of intraspecific variability in vertical habit use was observed. Four distinct depth distributions that were not related to tagging location, horizontal movements, sex, or size were detected. In addition, similar depth distributions did not necessitate similar dive patterns among sharks. Recognition of intraspecific variability in habitat use of top predators can be crucial for effective management of these species and for understanding their influence on ecosystem dynamics.     

Into the deep: the functionality of mesopelagic excursions by an oceanic apex predator

Comprehension of ecological processes in marine animals requires information regarding dynamic vertical habitat use. While many pelagic predators primarily associate with epipelagic waters, some species routinely dive beyond the deep scattering layer. Actuation for exploiting these aphotic habitats remains largely unknown. Recent telemetry data from oceanic whitetip sharks (Carcharhinus longimanus) in the Atlantic show a strong association with warm waters (>20°C) less than 200 m. Yet, individuals regularly exhibit excursions into the meso‐ and bathypelagic zone. In order to examine deep‐diving behavior in oceanic whitetip sharks, we physically recovered 16 pop‐up satellite archival tags and analyzed the high‐resolution depth and temperature data. Diving behavior was evaluated in the context of plausible functional behavior hypotheses including interactive behaviors, energy conservation, thermoregulation, navigation, and foraging. Mesopelagic excursions (n = 610) occurred throughout the entire migratory circuit in all individuals, with no indication of site specificity. Six depth‐versus‐time descent and ascent profiles were identified. Descent profile shapes showed little association with examined environmental variables. Contrastingly, ascent profile shapes were related to environmental factors and appear to represent unique behavioral responses to abiotic conditions present at the dive apex. However, environmental conditions may not be the sole factors influencing ascents, as ascent mode may be linked to intentional behaviors. While dive functionality remains unconfirmed, our study suggests that mesopelagic excursions relate to active foraging behavior or navigation. Dive timing, prey constituents, and dive shape support foraging as the most viable hypothesis for mesopelagic excursions, indicating that the oceanic whitetip shark may regularly survey extreme environments (deep depths, low temperatures) as a foraging strategy. At the apex of these deep‐water excursions, sharks exhibit a variable behavioral response, perhaps, indicating the presence or absence of prey.     

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.     

Women from Venus,men from Mars: inter‐sex foraging differences in the imperial cormorant Phalacrocorax atriceps a colonial seabird

Colonial seabirds are central place foragers and likely to be subject to substantial competition for resources. Mechanisms proposed for reducing intra‐specific competition include differential inter‐sex area use mediated by adult choice. We used GPS loggers and dive recorders to study area use and dive depth in a total of 27 male and 26 female imperial cormorants Phalacrocorax atriceps breeding at a colony of some 6500 birds at Punta Leon, Chubut, Argentina during 2004 and 2005. Although time spent travelling and distances between the colony and foraging sites were similar for both sexes, males and females travelled away from their colony using routes virtually perpendicular to each other so that their foraging areas were distinctly different; females hunted close to the coast while males foraged offshore in deeper water. Consideration of foraging efficiency underwater, defined as the duration spent on the bottom divided by the dive cycle duration, showed that females were more efficient at depths < 40 m while males more efficient at depths > 40 m. We suggest that the substantial sexual dimorphism in this species may be responsible for the different depth‐linked foraging efficiencies and that selection for appropriate depths could lead to differential habitat use and putative differences in prey selection.     

Vertical niche overlap by two ocean giants with similar diets: Ocean sunfish and leatherback turtles

We used satellite tags to record the patterns of depth utilisation for four ocean sunfish (Mola mola) and two leatherback turtles (Dermochelys coriacea) moving in broadly the same area off South Africa. Individuals were tracked for between 2 and 8 months and dive data relayed via satellite. For all the sunfish and one of the turtles, we received binned data on depth distribution, while for the second turtle we received individual dive profiles along with the proportion of time spent diving. Leatherback turtles dived almost exclusively within the upper 200 m, spending only 0.6 and 0.2% of their time > 200 m. There were times when sunfish likewise occupied these relatively shallow depths. However, there were also protracted periods when sunfish spent the majority of their time much deeper, with one individual remaining around 500 m for many hours at a time. These results suggest that sunfish sometimes exploit deeply distributed prey which is beyond the foraging range of leatherback turtles. We conclude that while both species are believed to feed predominantly on gelatinous zooplankton, the fact that sunfish do not need to come to the surface to breathe means that they can occupy an expanded vertical niche compared to the leatherback turtle.     

Sperm whale dive behavior characteristics derived from intermediate‐duration archival tag data

Here, we describe the diving behavior of sperm whales (Physeter macrocephalus) using the Advanced Dive Behavior (ADB) tag, which records depth data at 1‐Hz resolution and GPS‐quality locations for over 1 month, before releasing from the whale for recovery. A total of 27 ADB tags were deployed on sperm whales in the central Gulf of California, Mexico, during spring 2007 and 2008, of which 10 were recovered for data download. Tracking durations of all tags ranged from 0 to 34.5 days (median = 2.3 days), and 0.6 to 26.6 days (median = 5.0 days) for recovered tags. Recovered tags recorded a median of 50.8 GPS‐quality locations and 42.6 dives per day. Dive summary metrics were generated for archived dives and were subsequently classified into six categories using hierarchical cluster analysis. A mean of 77% of archived dives per individual were one of four dive categories with median Maximum Dive Depth >290 m (V‐shaped, Mid‐water, Benthic, or Variable), likely associated with foraging. Median Maximum Dive Depth was <30 m for the other two categories (Short‐ and Long‐duration shallow dives), likely representing socializing or resting behavior. Most tagged whales remained near the tagging area during the tracking period, but one moved north of Isla Tiburón, where it appeared to regularly dive to, and travel along the seafloor. Three whales were tagged on the same day in 2007 and subsequently traveled in close proximity (<1 km) for 2 days. During this period, the depth and timing of their dives were not coordinated, suggesting they were foraging on a vertically heterogeneous prey field. The multiweek dive records produced by ADB tags enabled us to generate a robust characterization of the diving behavior, activity budget, and individual variation for an important predator of the mesopelagos over temporal and spatial scales not previously possible.     

DIVE DATA FROM SATELLITE TAGS AND TIME-DEPTH RECORDERS: A COMPARISON IN WEDDELL SEAL PUPS

The diving behavior of juvenile Weddell seals, Leptonychotes weddellii , was monitored simultaneously with time-depth recorders (TDRs) and satellitelinked time-depth recorders (SLTDRs). Recovered TDRs provided a complete record of the depth and duration of all dives, while data received from SLTDR tags via the ARGOS satellite system were compressed into the number of dives in each of six depth or duration bins. The dive information from the two types of tags was compared to determine if data compression, processing, and transmission influenced the data received.
While only half of the dive data collected by TDRs was also received from the SLTDR tags, the chance of receiving SLTDR data was independent of when diving occurred, when data was transmitted, and the subsequent dive activity. In addition, the number of dives in each depth and duration bin was an accurate representation of the actual dive behavior. Therefore, SLTDR tags were judged to provide data qualitatively similar to that provided by TDRs. The accuracy of seal locations provided by Service ARGOS was estimated by comparison to Global Positioning System (GPS) locations, and the average position error found to be significantly greater than predicted by Service ARGOS or reported in other studies (LCO locations ± 11.4 km, LC1 ± 5.0 km).     

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.     

Short‐finned pilot whales (Globicephala macrorhynchus) of the Mariana Archipelago: Individual affiliations,movements, and spatial use

Little is known about short‐finned pilot whales (Globicephala macrorhynchus) in the western North Pacific outside of Japanese coastal waters. To expand understanding of short‐finned pilot whale ecology in the region, we conducted small‐boat surveys in 2010?2016 within the Mariana Archipelago to investigate individual associations, movements, spatial use, and dive behavior of short‐finned pilot whales. We collected genetic, photo‐identification, and satellite‐tag data and identified 191 distinctive individuals. A preliminary social network diagram of photo‐cataloged individuals revealed a main cluster that comprised 82% of individuals, representing all five mitochondrial DNA haplotypes identified within the population. Kernel density estimates for tagged short‐finned pilot whales (n = 11) during summer were used to identify areas with the highest probability of use (10% probability density contour), core area (50%) and home range (95%). The area with highest probability of use by short‐finned pilot whales was off the northwest side of Guam. Satellite tag data also suggest that some individuals are island‐associated year‐round. Data from five location‐dive tags demonstrated that the short‐finned pilot whales dove more often to intermediate depths at twilight and night, suggesting they may target prey that forage on the deep scattering layer as it migrates to and from the surface.     

Oxygen minimum zone: An important oceanographic habitat for deep‐diving northern elephant seals,Mirounga angustirostris

Little is known about the foraging behavior of top predators in the deep mesopelagic ocean. Elephant seals dive to the deep biota‐poor oxygen minimum zone (OMZ) (>800 m depth) despite high diving costs in terms of energy and time, but how they successfully forage in the OMZ remains largely unknown. Assessment of their feeding rate is the key to understanding their foraging behavior, but this has been challenging. Here, we assessed the feeding rate of 14 female northern elephant seals determined by jaw motion events (JME) and dive cycle time to examine how feeding rates varied with dive depth, particularly in the OMZ. We also obtained video footage from seal‐mounted videos to understand their feeding in the OMZ. While the diel vertical migration pattern was apparent for most depths of the JME, some very deep dives, beyond the normal diel depth ranges, occurred episodically during daylight hours. The midmesopelagic zone was the main foraging zone for all seals. Larger seals tended to show smaller numbers of JME and lower feeding rates than smaller seals during migration, suggesting that larger seals tended to feed on larger prey to satisfy their metabolic needs. Larger seals also dived frequently to the deep OMZ, possibly because of a greater diving ability than smaller seals, suggesting their dependency on food in the deeper depth zones. Video observations showed that seals encountered the rarely reported ragfish (Icosteus aenigmaticus) in the depths of the OMZ, which failed to show an escape response from the seals, suggesting that low oxygen concentrations might reduce prey mobility. Less mobile prey in OMZ would enhance the efficiency of foraging in this zone, especially for large seals that can dive deeper and longer. We suggest that the OMZ plays an important role in structuring the mesopelagic ecosystem and for the survival and evolution of elephant seals.     

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.     

Fish distributions along depth gradients of a sea mountain range conform to the mid‐domain effect

Species richness often peaks in the middle of bounded geographic domains (e.g. latitude, altitude or depth). Hump‐shaped richness distributions may be due to deterministic processes, such as adaptations to environmental variation. Alternatively, such distributions might also be due to stochastic process. The mid‐domain effect (MDE) posits that hump‐shaped richness distributions arise when species ranges are randomly arranged within the limits of the domain. We tested whether the MDE could account for the richness of bottom‐associated (demersal) fishes between 200 and 800 m on the Chatham Rise, New Zealand. We quantified the depth distributions of 59 fish species from 1891 research trawl catches made between 1991 and 2007. Results showed a broad plateau of high species richness near the centre of the domain (between 300 and 700 m), which was consistent with expectations of the MDE. Further, empirical species richness was better explained statistically by predictions of the MDE than models incorporating additional abiotic predictor variables. Our results deviated from previous studies that identified a greater richness of fishes in warmer, shallower depths with higher primary production. However, our study was conducted entirely below the euphotic zone, at depths where gradients are relatively weak, suggesting that support for the mid‐domain effect may increase across oceanic domains characterised by weak environmental gradients.     

First insights into the vertical habitat use of the whitespotted eagle ray Aetobatus narinari revealed by pop-up satellite archival tags

The whitespotted eagle ray Aetobatus narinari is a tropical to warm-temperate benthopelagic batoid that ranges widely throughout the western Atlantic Ocean. Despite conservation concerns for the species, its vertical habitat use and diving behaviour remain unknown. Patterns and drivers in the depth distribution of A. narinari were investigated at two separate locations, the western North Atlantic (Islands of Bermuda) and the eastern Gulf of Mexico (Sarasota, Florida, U.S.A.). Between 2010 and 2014, seven pop-up satellite archival tags were attached to A. narinari using three methods: a through-tail suture, an external tail-band and through-wing attachment. Retention time ranged from 0 to 180 days, with tags attached via the through-tail method retained longest. Tagged rays spent the majority of time (82.85 ± 12.17% S.D.) within the upper 10 m of the water column and, with one exception, no rays travelled deeper than ~26 m. One Bermuda ray recorded a maximum depth of 50.5 m, suggesting that these animals make excursions off the fore-reef slope of the Bermuda Platform. Individuals occupied deeper depths (7.42 ± 3.99 m S.D.) during the day versus night (4.90 ± 2.89 m S.D.), which may be explained by foraging and/or predator avoidance. Each individual experienced a significant difference in depth and temperature distributions over the diel cycle. There was evidence that mean hourly depth was best described by location and individual variation using a generalized additive mixed model approach. This is the first study to compare depth distributions of A. narinari from different locations and describe the thermal habitat for this species. Our study highlights the importance of region in describing A. narinari depth use, which may be relevant when developing management plans, whilst demonstrating that diel patterns appear to hold across individuals.     

Clinical assessment and postrelease monitoring of 11 mass stranded dolphins on Cape Cod,Massachusetts

The health, postrelease movements, and behavior of mass stranded Atlantic white‐sided dolphins (Lagenorhynchus acutus) and short‐beaked common dolphins (Delphinus delphis) from Cape Cod, Massachusetts, were evaluated. Health was assessed through physical examination and blood analysis. Eleven dolphins (eight white‐sided dolphins and three common dolphins) were relocated, outfitted with satellite transmitters, and released during seven mass stranding events. Five transmitters recorded only location, and six also included a time‐depth recorder. Transmission duration ranged from 8 h to 218 d, with a mean of 117 d (median = 118 d, SD = 82 d), after release. All dolphins demonstrated extensive movement throughout the Gulf of Maine. The distribution of tagged dolphins was considered normal based on comparisons with published data for these species. Excluding the dolphin that transmitted for only 8 h, mean minimum speeds for individual dolphins ranged from 3.4 to 6.6 km/h; overall mean for all dolphins was 5.4 km/h (SD = 0.9 km/h). The five dolphins with time‐depth recorders had mean dive depths of 8.6–40.3 m and mean dive durations of 46–296 s. Hematologic and biochemical data revealed only minor abnormalities. Data suggest that at least 10 of the 11 dolphins were likely successfully reintroduced into the wild.     

Using dive behavior and active acoustics to assess prey use and partitioning by fin and humpback whales near Kodiak Island,Alaska

Near the Kodiak Archipelago, fin (Balaenoptera physalus) and humpback (Megaptera novaeangliae) whales frequently overlap spatially and temporally. The Gulf Apex Predator‐prey study (GAP) investigated the prey use and potential prey partitioning between these sympatric species by combining concurrent analysis of vertical whale distribution with acoustic assessment of pelagic prey. Acoustic backscatter was classified as consistent with either fish or zooplankton. Whale dive depths were determined through suction cup tags. Tagged humpback whales (n = 10) were most often associated with distribution of fish, except when zooplankton density was very high. Associations between the dive depths of tagged fin whales (n = 4) and the vertical distribution of either prey type were less conclusive. However, prey assessment methods did not adequately describe the distribution of copepods, a potentially significant resource for fin whales. Mean dive parameters showed no significant difference between species when compared across all surveys. However, fin whales spent a greater proportion of dive time in the foraging phase than humpbacks, suggesting a possible difference in foraging efficiency between the two. These results suggest that humpback and fin whales may target different prey, with the greatest potential for diet overlap occurring when the density of zooplankton is very high.     

Ocean‐scale prediction of whale shark distribution

Aim Predicting distribution patterns of whale sharks (Rhincodon typus, Smith 1828) in the open ocean remains elusive owing to few pelagic records. We developed multivariate distribution models of seasonally variant whale shark distributions derived from tuna purse‐seine fishery data. We tested the hypotheses that whale sharks use a narrow temperature range, are more abundant in productive waters and select sites closer to continents than the open ocean. Location Indian Ocean. Methods We compared a 17‐year time series of observations of whale sharks associated with tuna purse‐seine sets with chlorophyll a concentration and sea surface temperature data extracted from satellite images. Different sets of pseudo‐absences based on random distributions, distance to shark locations and tuna catch were generated to account for spatiotemporal variation in sampling effort and probability of detection. We applied generalized linear, spatial mixed‐effects and Maximum Entropy models to predict seasonal variation in habitat suitability and produced maps of distribution. Results The saturated generalized linear models including bathymetric slope, depth, distance to shore, the quadratic of mean sea surface temperature, sea surface temperature variance and chlorophyll a had the highest relative statistical support, with the highest percent deviance explained when using random pseudo‐absences with fixed effect‐only models and the tuna pseudo‐absences with mixed‐effects models (e.g. 58% and 26% in autumn, respectively). Maximum Entropy results suggested that whale sharks responded mainly to variation in depth, chlorophyll a and temperature in all seasons. Bathymetric slope had only a minor influence on the presence. Main conclusions Whale shark habitat suitability in the Indian Ocean is mainly correlated with spatial variation in sea surface temperature. The relative influence of this predictor provides a basis for predicting habitat suitability in the open ocean, possibly giving insights into the migratory behaviour of the world’s largest fish. Our results also provide a baseline for temperature‐dependent predictions of distributional changes in the future.     

Summer diving and haul‐out behavior of leopard seals (Hydrurga leptonyx) near mesopredator breeding colonies at Livingston Island,Antarctic Peninsula

Leopard seals are conspicuous apex predators in Antarctic coastal ecosystems, yet their foraging ecology is poorly understood. Historically, the ecology of diving vertebrates has been studied using high‐resolution time‐depth records; however, to date such data have not been available for leopard seals. Twenty‐one time‐depth recorders were deployed on seasonally resident adult females in January and February between 2008 and 2014. The average deployment length was 13.65 ± 11.45 d and 40,308 postfilter dives were recorded on 229 foraging trips. Dive durations averaged 2.20 ± 1.23 min. Dives were shallow with 90.1% measuring 30 m or less, and a mean maximum dive depth of 16.60 ± 10.99 m. Four dive types were classified using a k‐means cluster analysis and compared with corresponding animal‐borne video data. Dive activity (number of dives/hour) was concentrated at night, including crepuscular periods. Haul‐out probabilities were highest near midday and were positively correlated with available daylight. Visual observations and comparisons of diving activity between and within years suggest individual‐based differences of foraging effort by time of day. Finally, dive and video data indicate that in addition to at‐surface hunting, benthic searching and facultative scavenging are important foraging strategies for leopard seals near coastal mesopredator breeding colonies.     

Red muscle proportions and enzyme activities in deep‐sea demersal fishes

Owing to the paucity of data on the red muscle of deep‐sea fishes, the goal of this study was to determine the proportions of red muscle in demersal fishes and its enzymatic activities to characterize how routine swimming abilities change with depths of occurrence. Cross sectional analysis of the trunk musculature was used to evaluate the proportion of red muscle in 38 species of Californian demersal fishes living at depths between 100 and 3000 m. The activity of metabolic enzymes was also assayed in a sub‐set of 18 species. Benthic fishes had lower proportions of red muscle and lower metabolic enzyme activities than benthopelagic species. Mean proportion of red muscle declined significantly with depth with the greatest range of values in shallow waters and species with low proportions found at all depths. This suggested that while sedentary species occur at all depths, the most active species occur in shallow waters. Citrate synthase activity declined significantly with depth across all species, indicating that the mass‐specific metabolic capacity of red muscle is lower in deep‐sea species. These patterns may be explained by coupling of red and white muscle physiologies, a decrease in physical energy of the environment with depth or by the prevalence of anguilliform body forms and swimming modes in deep‐living species.     

Habitat use and movement patterns of bull sharks Carcharhinus leucas determined using pop‐up satellite archival tags

Habitat use, movement and residency of bull sharks Carcharhinus leucas were determined using satellite pop‐up archival transmitting (PAT) tags throughout coastal areas in the U.S., Gulf of Mexico and waters off the south‐east U.S. From 2005 to 2007, 18 fish (mean size = 164 cm fork length, L_F) were tagged over all seasons. Fish retained tags for up to 85 days (median = 30 days). Based on geolocation data from initial tagging location to pop‐off location, C. leucas generally travelled c. 5–6 km day^?1 and travelled an average of 143·6 km. Overall, mean proportions of time at depth revealed C. leucas spent the majority of their time in waters <20 m. They exhibited significant differences among depths but were not found at a particular depth regardless of diurnal period. Most fish occupied temperatures c. 32° C with individuals found mostly between 26 and 33° C. Geolocation data for C. leucas were generally poor and varied considerably but tracks for two individuals revealed long distance movements. One fish travelled from the south‐east coast of the U.S. to coastal Texas near Galveston while another moved up the east coast of the U.S. to South Carolina. Data on C. leucas movements indicated that they are found primarily in shallower waters and tend to remain in the same location over long periods. While some individuals made large‐scale movements over open ocean areas, the results emphasize the importance of the coastal zone for this species as potential essential habitat, particularly in areas of high freshwater inflow.