Temporal patterns in the acoustic signals of beaked whales at Cross Seamount

Seamounts may influence the distribution of marine mammals through a combination of increased ocean mixing, enhanced local productivity and greater prey availability. To study the effects of seamounts on the presence and acoustic behaviour of cetaceans, we deployed a high-frequency acoustic recording package on the summit of Cross Seamount during April through October 2005. The most frequently detected cetacean vocalizations were echolocation sounds similar to those produced by ziphiid and mesoplodont beaked whales together with buzz-type signals consistent with prey-capture attempts. Beaked whale signals occurred almost entirely at night throughout the six-month deployment. Measurements of prey presence with a Simrad EK-60 fisheries acoustics echo sounder indicate that Cross Seamount may enhance local productivity in near-surface waters. Concentrations of micronekton were aggregated over the seamount in near-surface waters at night, and dense concentrations of nekton were detected across the surface of the summit. Our results suggest that seamounts may provide enhanced foraging opportunities for beaked whales during the night through a combination of increased productivity, vertical migrations by micronekton and local retention of prey. Furthermore, the summit of the seamount may act as a barrier against which whales concentrate prey.     

HOW DO SPERM WHALES CATCH SQUIDS?

Vision may play a central role in sperm whale predation. Two complementary hypotheses regarding the detection and capture of prey items are presented, based on a review of mesopelagic ecology. The first hypothesis postulates that sperm whales locate their prey visually, either silhouetted against the midwater "sky," or by searching for bioluminescence produced by the movements of their prey. The second hypothesis postulates that sperm whales create a zone of stimulated bioluminescence around the mouth, which attracts squids and other visual predators. Studies of midwater fishes and invertebrates document the importance of vision in mesopelagic communities. If sperm whales search for silhouetted prey, they should be oriented upside-down to improve visual coverage and to facilitate the transition from search to prey capture. Prey capture events should be marked by excursions toward the surface. If they lure their prey, they should swim at a steady pace, with little rapid acceleration, and spend most of their time foraging at depths with the greatest potential for stimulated bioluminescence.     

Shallow food for deep divers: Dynamic foraging behavior of male sperm whales in a high latitude habitat

Groups of female and immature sperm whales live at low latitudes and show a stereotypical diving and foraging behavior with dives lasting about 45 min to depths of between 400 and 1200 m. In comparison, physically mature male sperm whales migrate to high latitudes where little is known about their foraging behavior and ecology. Here we use acoustic recording tags to study the diving and acoustic behavior of male sperm whales foraging off northern Norway. Sixty-five hours of tag data provide detailed information about the movements and sound repertoire of four male sperm whales performing 83 dives lasting between 6 and 60 min. Dives ranged in depth between 14 and 1860 m, with a median depth of 175 m, and 92% of the surfacings lasted less than 15 min. The four whales clicked for an average 91% (SD = 10) of the dive duration, where the first usual click was produced at depths ranging between 4 and 218 m and the last usual click at depths ranging between 1 and 1114 m. Echolocation buzzes, which are used as an indication of prey capture attempts, were emitted at depths between 17 and 1860 m, during both the descent and ascent phase of deep dives. The foraging behavior varied markedly with depth, with the timing and duration of prey capture attempts during shallow dives suggesting that the whales target more sparsely distributed prey. In contrast, deep dives involve frequent prey capture attempts and seem to target more dense food layers. The evidence of exploitation of different food layers, including epipelagic prey, is consistent with the hypothesis that male sperm whales may migrate to high latitudes to access a productive, multi-layered foraging habitat.     

CHARACTERIZATION OF BEAKED WHALE (ZIPHIIDAE) AND SPERM WHALE (PHYSETER MACROCEPHALUS) SUMMER HABITAT IN SHELF-EDGE AND DEEPER WATERS OFF THE NORTHEAST U. S.

Sperm whales ( Physeter macrocephalus ) and beaked whales ( Mesoplodon spp. and Ziphius cavirostris ) are deep-diving cetaceans that frequent shelf-edge and Gulf Stream waters off the northeast U. S. coast. Sighting data collected during seven summer (1990, 1991, 1993, and 1995–1998) shipboard surveys were analyzed using a geographic information system to determine habitat use based on bathymetric and oceanographic features. Although sighting rates were lower for beaked whales, both taxa occupied similar habitats. Beaked whales were concentrated at the colder shelf edge, whereas sperm whales were associated with warmer off-shelf water. Mean sighting rates for both taxa were higher in canyon features, but only beaked whale sighting rates were significantly different between canyon and non-canyon habitat (Wilcoxon signed rank test P = 0.007). Within the shared habitat, the two taxa were separated at fine-scale based on oceanographic features.     

Building time-budgets from bioacoustic signals to measure population-level changes in behavior: a case study with sperm whales in the Gulf of Mexico

Evaluating changes in the collective behavior of a population can be an indirect method for inferring organismal responses to changing environmental conditions. Apex predators, such as the sperm whale (Physeter macrocephalus), can provide valuable insights into the ecosystem processes of the deep sea, where little direct observation can be made. Sperm whales are often difficult to observe at sea, as they inhabit deep, offshore waters and spend most of their lives beneath the surface. However, sperm whales are extremely amenable to passive acoustic monitoring, as their vocalizations are well-studied, highly distinguishable, produced regularly, and can be detected at relatively long ranges (>10 km). Sperm whales produce distinct clicks in two behavioral contexts (social interaction or foraging/prey capture); thus, we can use acoustic detection of these vocalizations to infer patterns of large-scale, collective behavior, which is similar to studying calling frogs or insects indicating their reproductive phenology. We recorded behaviorally-specific sperm whale vocalizations at three sites in the Northern Gulf of Mexico in July 2010 and 2011. We used these recordings to construct population-level time budgets, an empirical collective metric of behavior, based on the ratio of hours in a day with social clicks to the hours in a day with foraging clicks, and represented this as an “acoustic activity index.” Our index showed significant differences in the proportions of social and foraging behavior across the range of sperm whales in the Northern Gulf of Mexico, and the proportion of social activity increased by more than a factor of two from 2010 to 2011. These differences support previous evidence of differential habitat use by sperm whales in the Gulf of Mexico, and suggest possible changes in environmental conditions between years. Thus, the acoustic activity index may provide a powerful way to evaluate changes in behavior and link them to changing ecological conditions. This novel application of bioacoustics to constructing time budgets and creating a behaviorally-based index at the population scale can serve as an indicator of ecological change, and greatly enhance our ability to understand the behavior and ecology of many acoustically active species.     

CONSEQUENCES OF ALTERNATIVE FUNCTIONAL RESPONSE FORMULATIONS IN MODELS EXPLORING WHALE-FISHERY INTERACTIONS

We evaluated the utility of Ecosim for exploring interactions between cetacean predators, their prey, and fisheries. We formulated six Ecosim parameterizations, representing alternative hypotheses of feeding interactions (functional response) between cetaceans and their main fish prey, and examined differences in the predicted responses to simulated harvesting regimes for minke whales and their prey. Regardless of the type of function response formulated, intense fishing on the main fish prey of minke whales had a longer-lasting negative impact on minke whales than when minke whale biomass was removed directly by harvesting. Consumption rate, biomass, feeding time and mortality of minke whales were all sensitive to the type of functional response specified. Inclusion of "handling time" limited minke whales consumption at high prey densities and predicted higher consumption at low prey densities; features characteristic of a type II functional response. Predicted decline and recovery rates of minke whales were slower than when consumption rates were not limited. Addition of "foraging time" adjustments resulted in more conservative estimates of decline and recovery. However, when "other mortality" was linked to time spent foraging, exposure to higher mortality at low prey densities, and reduced mortality at high prey densities resulted in dramatic differences in predicted biomass trajectory. Sensitivity to the "other mortality" assumption is important for cetaceans whose predation mortality is only a small proportion of total mortality. Differences in the feeding and biomass dynamics were also observed when prey availability to predators was represented by changes in prey vulnerability, confirming earlier reports that Ecosim predictions are sensitive to this parameter.     

Odontocete occurrence in relation to changes in oceanography at a remote equatorial Pacific seamount

Seamounts are considered hot spots of biodiversity and can aggregate pelagic predators and their prey. Passive acoustic monitoring was conducted over 3 mo in 2012 to document the occurrence of odontocetes near a seamount chain in the central equatorial Pacific in relation to oceanographic changes over time. Beaked whale echolocation signals were most frequently encountered. The main beaked whale signal was an unknown type, BW38, which resembled signals produced by Blainville's beaked whales. It had high occurrence during high sea surface temperature and low sea surface salinity. Cuvier's beaked whales were the second most detected. They had an opposite pattern and were encountered more often when sea surface temperature was low and net primary productivity was high. Risso's dolphins and short‐finned pilot whales had high acoustic densities, and echolocated predominantly at night. Risso's dolphins occurred more often during low sea surface height deviation. False killer whales were less frequently detected and mostly occurred during the day. Sperm whale detections were fewer than expected and associated with high chlorophyll a. Short duration Kogiidae encounters occurred on average every third day. These types of long‐term site studies are an informative tool to comparatively assess species composition, relative abundance, and relationship to oceanographic changes.     

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

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

Predicting top predator habitats in the Southwest Indian Ocean

Top predators need to develop optimal strategies of resources and habitats utilization in order to optimize their foraging success. At the individual scale, a predator has to maximize his intake of food while minimizing his cost of foraging to optimize his energetic gain. At the ecosystem scale, we hypothesized that foraging strategies of predators also respond to their general energetic constraints. Predators with energetically costly lifestyles may be constrained to select high quality habitats whereas more phlegmatic predators may occupy both low and high quality habitats. The objectives of this study were 1) to investigate predator responses to heterogeneity in habitat quality with reference to their energetic strategies and 2) to evaluate their responses to contemporaneous versus averaged habitat quality. We collected cetacean and seabird data from an aerial survey in the Southwest Indian Ocean, a region characterized by heterogeneous oceanographic conditions. We classified cetaceans and seabirds into energetic guilds and described their habitats using remotely sensed covariates at contemporaneous and time‐averaged resolutions and static covariates. We used generalized additive models to predict their habitats at the regional scale. Strategies of habitat utilization appeared in accordance with predators energetic constraints. Cetaceans responded to the heterogeneity in habitat quality, with higher densities predicted in more productive areas. However, the costly Delphininae appeared to be more dependent on habitat quality (showing a 1‐to‐13 ratio between the lowest and highest density sectors) than the more phlegmatic sperm and beaked whales (showing only a 1‐to‐3 ratio). For seabirds, predictions primarily reflected colony locations, although the colony effect was stronger for costly seabirds. Moreover, our results suggest that predators may respond better to persistent oceanographic features. To provide a third dimension to habitat quality, cetacean strategies of utilization of the vertical habitat could be related to the distribution of micronekton in the water column.     

The effects of prey demography on humpback whale (Megaptera novaeangliae) abundance around Anvers Island, Antarctica

Baleen whales and Adelie penguins in the near-shore waters around the Antarctic Peninsula forage principally on Antarctic krill. Given the spatial overlap in the distribution of these krill predators (particularly humpback whales) and their dependence on krill, the goals of this paper are to determine if the inter-annual community structure and relative abundance of baleen whales around Anvers Island is related to krill demography and abundance, and if the potential exists for inter-specific interactions between Adelie penguins and baleen. We use whale sightings and prey data from both net tows and Adelie penguin stomach samples to correlate the abundance of humpback whales with krill demography and abundance from 1993 to 2001. We find significant relationships between whale abundance and the size–frequency distribution of krill targeted by Adelie penguins, as well as the foraging success of Adelie penguins. These findings suggest both krill predators share common prey preferences in the upper portions of the water column around Anvers Island. These findings highlight the need for better knowledge of baleen whale foraging ecology and inter-specific interactions with penguins, as sea ice and krill populations around the Antarctic Peninsula are affected by rapid changes in climate.     

Linking killer whale survival and prey abundance: food limitation in the oceans' apex predator?

Killer whales (Orcinus orca) are large predators that occupy the top trophic position in the world''s oceans and as such may have important roles in marine ecosystem dynamics. Although the possible top-down effects of killer whale predation on populations of their prey have received much recent attention, little is known of how the abundance of these predators may be limited by bottom-up processes. Here we show, using 25 years of demographic data from two populations of fish-eating killer whales in the northeastern Pacific Ocean, that population trends are driven largely by changes in survival, and that survival rates are strongly correlated with the availability of their principal prey species, Chinook salmon (Oncorhynchus tshawytscha). Our results suggest that, although these killer whales may consume a variety of fish species, they are highly specialized and dependent on this single salmonid species to an extent that it is a limiting factor in their population dynamics. Other ecologically specialized killer whale populations may be similarly constrained to a narrow range of prey species by culturally inherited foraging strategies, and thus are limited in their ability to adapt rapidly to changing prey availability.     

Sperm whale behaviour indicates the use of echolocation click buzzes "creaks" in prey capture

During foraging dives, sperm whales (Physeter macrocephalus) produce long series of regular clicks at 0.5-2 s intervals interspersed with rapid-click buzzes called "creaks". Sound, depth and orientation recording Dtags were attached to 23 whales in the Ligurian Sea and Gulf of Mexico to test whether the behaviour of diving sperm whales supports the hypothesis that creaks are produced during prey capture. Sperm whales spent most of their bottom time within one or two depth bands, apparently feeding in vertically stratified prey layers. Creak rates were highest during the bottom phase: 99.8% of creaks were produced in the deepest 50% of dives, 57% in the deepest 15% of dives. Whales swam actively during the bottom phase, producing a mean of 12.5 depth inflections per dive. A mean of 32% of creaks produced during the bottom phase occurred within 10 s of an inflection (13x more than chance). Sperm whales actively altered their body orientation throughout the bottom phase with significantly increased rates of change during creaks, reflecting increased manoeuvring. Sperm whales increased their bottom foraging time when creak rates were higher. These results all strongly support the hypothesis that creaks are an echolocation signal adapted for foraging, analogous to terminal buzzes in taxonomically diverse echolocating species.     

Oceanic thermal structure mediates dive sequences in a foraging seabird

1. Changes in marine ecosystems are easier to detect in upper‐level predators, like seabirds, which integrate trophic interactions throughout the food web.
2. Here, we examined whether diving parameters and complexity in the temporal organization of diving behavior of little penguins (Eudyptula minor) are influenced by sea surface temperature (SST), water stratification, and wind speed—three oceanographic features influencing prey abundance and distribution in the water column.
3. Using fractal time series analysis, we found that foraging complexity, expressed as the degree of long‐range correlations or memory in the dive series, was associated with SST and water stratification throughout the breeding season, but not with wind speed. Little penguins foraging in warmer/more‐stratified waters exhibited greater determinism (memory) in foraging sequences, likely as a response to prey aggregations near the thermocline. They also showed higher foraging efficiency, performed more dives and dove to shallower depths than those foraging in colder/less‐stratified waters.
4. Reductions in the long‐term memory of dive sequences, or in other words increases in behavioral stochasticity, may suggest different strategies concerning the exploration–exploitation trade‐off under contrasting environmental conditions.

     

Using Models of Social Transmission to Examine the Spread of Longline Depredation Behavior among Sperm Whales in the Gulf of Alaska

Fishing, farming and ranching provide opportunities for predators to prey on resources concentrated by humans, a behavior termed depredation. In the Gulf of Alaska, observations of sperm whales depredating on fish caught on demersal longline gear dates back to the 1970s, with reported incidents increasing in the mid-1990s. Sperm whale depredation provides an opportunity to study the spread of a novel foraging behavior within a population. Data were collected during National Marine Fisheries Service longline surveys using demersal longline gear in waters off Alaska from 1998 to 2010. We evaluated whether observations of depredation fit predictions of social transmission by fitting the temporal and spatial spread of new observations of depredation to the Wave of Advance model. We found a significant, positive relationship between time and the distance of new observations from the diffusion center (r² = 0.55, p-value  = 0.003). The data provide circumstantial evidence for social transmission of depredation. We discuss how changes in human activities in the region (fishing methods and regulations) have created a situation in which there is spatial-temporal overlap with foraging sperm whales, likely influencing when and how the behavior spread among the population.     

Following a foraging fish-finder: diel habitat use of Blainville's beaked whales revealed by echolocation

Simultaneous high resolution sampling of predator behavior and habitat characteristics is often difficult to achieve despite its importance in understanding the foraging decisions and habitat use of predators. Here we tap into the biosonar system of Blainville''s beaked whales, Mesoplodon densirostris, using sound and orientation recording tags to uncover prey-finding cues available to echolocating predators in the deep-sea. Echolocation sounds indicate where whales search and encounter prey, as well as the altitude of whales above the sea-floor and the density of organisms around them, providing a link between foraging activity and the bio-physical environment. Tagged whales (n = 9) hunted exclusively at depth, investing most of their search time either in the lower part of the deep scattering layer (DSL) or near the sea-floor with little diel change. At least 43% (420/974) of recorded prey-capture attempts were performed within the benthic boundary layer despite a wide range of dive depths, and many dives included both meso- and bentho-pelagic foraging. Blainville''s beaked whales only initiate searching when already deep in the descent and encounter prey suitable for capture within 2 min of the start of echolocation, suggesting that these whales are accessing prey in reliable vertical strata. Moreover, these prey resources are sufficiently dense to feed the animals in what is effectively four hours of hunting per day enabling a strategy in which long dives to exploit numerous deep-prey with low nutritional value require protracted recovery periods (average 1.5 h) between dives. This apparent searching efficiency maybe aided by inhabiting steep undersea slopes with access to both the DSL and the sea-floor over small spatial scales. Aggregations of prey in these biotopes are located using biosonar-derived landmarks and represent stable and abundant resources for Blainville''s beaked whales in the otherwise food-limited deep-ocean.     

Partitioning mechanisms of Predator Interference in different Habitats

Prey are often consumed by multiple predator species. Predation rates on shared prey species measured in isolation often do not combine additively due to interference or facilitation among the predator species. Furthermore, the strength of predator interactions and resulting prey mortality may change with habitat type. We experimentally examined predation on amphipods in rock and algal habitats by two species of intertidal crabs, Hemigrapsus sanguineus (top predators) and Carcinus maenas (intermediate predators). Algae provided a safer habitat for amphipods when they were exposed to only a single predator species. When both predator species were present, mortality of amphipods was less than additive in both habitats. However, amphipod mortality was reduced more in rock than algal habitat because intermediate predators were less protected in rock habitat and were increasingly targeted by omnivorous top predators. We found that prey mortality in general was reduced by (1) altered foraging behavior of intermediate predators in the presence of top predators, (2) top predators switching to foraging on intermediate predators rather than shared prey, and (3) density reduction of intermediate predators. The relative importance of these three mechanisms was the same in both habitats; however, the magnitude of each was greater in rock habitat. Our study demonstrates that the strength of specific mechanisms of interference between top and intermediate predators can be quantified but cautions that these results may be habitat specific. An erratum to this article can be found at     

Movements of foraging king penguins through marine mesoscale eddies

Despite increasing evidence that marine predators associate with mesoscale eddies, how these marine features influence foraging movements is still unclear. This study investigates the relationship of at-sea movements of king penguins to mesoscale eddies using oceanographic remote sensing and movement data from 43 individual trips over 4 years. Simultaneous satellite measurements provided information on gradients of sea surface temperature and currents associated with eddies determined from altimetry. Penguins tended to swim rapidly with currents as they travelled towards foraging zones. Swimming speed indicative of foraging occurred within mesoscale fronts and strong currents associated with eddies at the Polar Front. These results demonstrate the importance of mesoscale eddies in directing foraging efforts to allow predators to rapidly get to rich areas where high concentrations of prey are likely to be encountered. When returning to the colony to relieve the incubating partner or to feed the chick, the birds followed a direct and rapid path, seemingly ignoring currents.     

DISTRIBUTION OF KILLER WHALE PODS IN PRINCE WILLIAM SOUND, ALASKA 1984–1996

Thirteen years of encounter data (1984–1996) were used to examine killer whale distribution within Prince William Sound, Alaska. Four patterns of area use were found, which comprised differences between resident pods and transient groups and differences among resident groups. Resident pods frequented large open passages, while transient groups used the narrow passages and bays in the southwest. This dichotomy likely reflects resident use of salmon and transient use of pinniped prey resources, as well as the different foraging strategies required for these prey types. Four resident pods (AB, AI, AJ, and AN) used Knight Island Passage more than other areas of the Sound; two (AE and AK) used all areas of the Sound more evenly. Use of the Sound by the AT1 transient whales declined in the latter part of the study. Nearshore foraging for pinniped prey by the AT1 transient whales was more common in areas where these whales spend a disproportionate amount of time, suggesting that these areas were critical foraging habitat for them. No similar pattern emerged for Open-water Foraging for cetaceans by AT1 whales, nor for foraging by the resident whales.     

Combining Methods to Describe Important Marine Habitats for Top Predators: Application to Identify Biological Hotspots in Tropical Waters

In tropical waters resources are usually scarce and patchy, and predatory species generally show specific adaptations for foraging. Tropical seabirds often forage in association with sub-surface predators that create feeding opportunities by bringing prey close to the surface, and the birds often aggregate in large multispecific flocks. Here we hypothesize that frigatebirds, a tropical seabird adapted to foraging with low energetic costs, could be a good predictor of the distribution of their associated predatory species, including other seabirds (e.g. boobies, terns) and subsurface predators (e.g., dolphins, tunas). To test this hypothesis, we compared distribution patterns of marine predators in the Mozambique Channel based on a long-term dataset of both vessel- and aerial surveys, as well as tracking data of frigatebirds. By developing species distribution models (SDMs), we identified key marine areas for tropical predators in relation to contemporaneous oceanographic features to investigate multi-species spatial overlap areas and identify predator hotspots in the Mozambique Channel. SDMs reasonably matched observed patterns and both static (e.g. bathymetry) and dynamic (e.g. Chlorophyll a concentration and sea surface temperature) factors were important explaining predator distribution patterns. We found that the distribution of frigatebirds included the distributions of the associated species. The central part of the channel appeared to be the best habitat for the four groups of species considered in this study (frigatebirds, brown terns, boobies and sub-surface predators).     

Migration pathways, behavioural thermoregulation and overwintering grounds of blue sharks in the Northwest Atlantic

The blue shark Prionace glauca is the most abundant large pelagic shark in the Atlantic Ocean. Although recaptures of tagged sharks have shown that the species is highly migratory, migration pathways towards the overwintering grounds remain poorly understood. We used archival satellite pop-up tags to track 23 blue sharks over a mean period of 88 days as they departed the coastal waters of North America in the autumn. Within 1-2 days of entering the Gulf Stream (median date of 21 Oct), all sharks initiated a striking diel vertical migration, taking them from a mean nighttime depth of 74 m to a mean depth of 412 m during the day as they appeared to pursue vertically migrating squid and fish prey. Although functionally blind at depth, calculations suggest that there would be a ~2.5-fold thermoregulatory advantage to swimming and feeding in the markedly cooler deep waters, even if there was any reduced foraging success associated with the extreme depth. Noting that the Gulf Stream current speeds are reduced at depth, we used a detailed circulation model of the North Atlantic to examine the influence of the diving behaviour on the advection experienced by the sharks. However, there was no indication that the shark diving resulted in a significant modification of their net migratory pathway. The relative abundance of deep-diving sharks, swordfish, and sperm whales in the Gulf Stream and adjacent waters suggests that it may serve as a key winter feeding ground for large pelagic predators in the North Atlantic.