Seasonal changes in the diving parameters of king penguins (Aptenodytes patagonicus)

Contrasting conditions at-sea are likely to affect the foraging behaviour of seabirds. However, the effect of season on the dive parameters of penguins is poorly known. We report here on an extensive study of the diving behaviour of king penguins (Aptenodytes patagonicus) over the bird's complete annual cycle at the Crozet Islands. Time-depth recorders were used to record dive duration, bottom duration, post-dive interval, ascent rate and descent rate in breeding adults during different seasons in 1995 and 1996. Seasons included summer (n=6, incubation; n=6, chick brooding), autumn and winter (n=5 and n=3, respectively, chick at the crèche stage), and spring (n=4, birds at the post-moult stage). In all seasons dive duration increased with dive depth, but, for a given depth, dives were longer in winter (6.8 min when averaged over the 100-210 m depth layer) than in spring (4.6 min) and summer (4.4 min). The time spent at the bottom of the dives, which probably represents a substantial part of the feeding time, was much longer in winter (2.5 min per dive for dives over the 100-210 m layer) than during other seasons (1.0-1.4 min), i.e. there was a 2.5-fold augmentation for similar diving depths. Ascent and descent rates increased with increasing dive depth, but no difference in the relationships between rates of ascent and descent and dive depth was found among seasons. Furthermore, for all dive depths, ascent and descent rates were independent of the bottom duration. In all seasons post-dive intervals increased with dive duration and with dive depth, but they were longer in spring (2.3 min for dives over the 100-210 m layer) and summer than in autumn and winter (1.6-1.8 min). The diving efficiency decreased with increasing dive depth and was higher in autumn and winter (0.22-0.29) than in summer and spring (0.15-0.18). The large increase in bottom and dive duration from spring to winter is in agreement with the seasonal drop in prey density, with penguins spending more time searching for prey. In contrast, the consistency of the vertical velocity during contrasting conditions at-sea suggests that the transit time to depth is an important component of the foraging behaviour (scanning of the water column) that is independent of the prey availability. The time budget of the penguins during diving in a fluctuating environment appears to vary primarily during the bottom phase of the dives, with bottom duration increasing with diminishing prey supplies, while post-dive intervals shorten in the same time.     

GPS and time-depth loggers reveal underwater foraging plasticity in a flying diver,the Cape Cormorant

Knowledge on how divers exploit the water column vertically in relation to water depth is crucial to our understanding of their ecology and to their subsequent conservation. However, information is still lacking for the smaller-bodied species, due mostly to size constraints of data-loggers. Here, we report the diving behaviour of a flying diving seabird, the Cape Cormorant Phalacrocorax capensis, weighing 1.0–1.4 kg. Results were obtained by simultaneously deploying small, high resolution and high sampling frequency GPS and time-depth loggers on birds breeding on islands off Western South Africa (34°S, 18°E) in 2008. In all, dive category was assigned to all dives performed by 29 birds. Pelagic dives occurred almost as frequently as benthic dives. Pelagic dives were shallow (mean: 5 m) and took place over seafloors 5–100 m deep. Benthic dives were deeper, occurring on seafloors mainly 10–30 m deep. Dive shape was linked to dive category in only 60% of dives, while the descent rate, ascent rate and bottom duration/dive duration ratio of a dive best explained its dive category. This shows that only the concomitant use of tracking and depth tags can adequately classify diving strategies in a diver like the Cape Cormorant. Diet was mainly Cape Anchovy Engraulis encrasicolis, suggesting that birds probably displayed two contrasted strategies for capturing the same prey. Flexible foraging techniques represent an important key to survival inside the highly productive but heterogeneous Benguela upwelling ecosystem.     

Foraging strategies and prey encounter rate of free-ranging Little Penguins

There is little information on the effort put into foraging by seabirds, even though it is fundamental to many issues in behavioural ecology. Recent researchers have used changes in the underwater cruising speed of penguins to allude to prey ingestion since accelerations are thought to reflect the encounter and pursuit of prey. In this study, we attached minute accelerometers, to determine flipper beat frequency as a proxy for prey pursuit, to Little Penguins Eudyptula minor foraging in shallow waters in Western Australia. During diving, Little Penguins flapped continuously and at a regular pace of 3.16 Hz while descending the water column and throughout the bottom phase of most dives. However, the frequency and amplitude of wingbeats increased transitorily, reaching 3.5–5.5 Hz, during some dives indicating prey pursuit. Pursuit phases lasted a mean of 2.9±3.3 s and occurred principally during the bottom phases of dives (75.4%). Most dives in all birds (86%) had a clear square-shaped depth profile indicating feeding activity near the seabed in the shallow waters of the bays. Hourly maximum depth, time spent underwater, percentage of dives with pursuit events and catch per unit effort showed an overall increase from zero at ca. 0500 h to a maximum during the hours around mid-day before decreasing to zero by 1900 h. During pursuit phases, Little Penguins headed predominantly downward, probably using the seabed to assist them in trapping their prey. In the light of our results, we discuss depth use by Little Penguins and their allocation of foraging effort and prey capture success as a function of environmental conditions.     

Diving behaviour of female northern rockhopper penguins, Eudyptes chrysocome moseleyi, during the brooding period at Amsterdam Island (Southern Indian Ocean)

The pattern and characteristics of diving in 14 female northern rockhopper penguins, Eudyptes chrysocome moseleyi, were studied at Amsterdam Island (37°50′S; 77°31′E) during the guard stage, using electronic time–depth recorders. Twenty-nine foraging trips (27 daily foraging trips and two longer trips including one night) with a total of 16 572 dives of ≥3 m were recorded. Females typically left the colony at dawn and returned in the late afternoon, spending an average of 12 h at sea, during which they performed ∼550 dives. They were essentially inshore foragers (mean estimated foraging range 6 km), and mainly preyed upon the pelagic euphausiid Thysanoessa gregaria, fishes and squid being only minor components of the diet. Mean dive depth, dive duration, and post-dive intervals were 18.4 m (max. depth 109 m), 57 s (max. dive duration 168 s), and 21 s (37% of dive duration), respectively. Descent and ascent rates averaged 1.2 and 1.0 ms⁻¹ and were, together with dive duration, significantly correlated with dive depth. Birds spent 18% of their total diving time in dives reaching 15 to 20 m, and the mean maximum diving efficiency (bottom time:dive cycle duration) occurred for dives reaching 15 to 35 m. The most remarkable feature of diving behaviour in northern rockhopper penguins was the high percentage of time spent diving during daily foraging trips (on average, 69% of their time at sea); this was mainly due to a high dive frequency (∼44 dives per hour), which explained the high total vertical distance travelled during one trip (18 km on average). Diving activity at night was greatly reduced, suggesting that, as other penguins, E. chrysocome moseleyi are essentially diurnal, and locate prey using visual cues. Received: 9 December 1998 / Accepted: 3 March 1999     

Swim speeds of free-ranging great cormorants

Information about foraging speeds is particularly valuable when the impact of a predator species upon a community of prey has to be defined, as in the case of great cormorants. We measured the swim speed of 12 (six males and six females) free-ranging great cormorants Phalacrocorax carbo, foraging off the Greenland coast during the summer of 2003, using miniaturized data-loggers. Although mean body mass of males was 27% greater than that of females, and mean swim speed of males were 29–57% higher than that of females during foraging phases (but not descent phases) of dives, these differences in speeds were not significant due to high variances. Birds descended to the mean maximum depth of 4.7 m at an average speed of 1.6±0.5 m s⁻¹, a speed similar to that measured in captive cormorants in previous studies. Although bursts of up to 4 m s⁻¹ were recorded, speed usually decreased during the deepest (foraging) phase of dives, being on average 0.8±0.6 m s⁻¹. Speeds measured here should be taken with caution, because the large propeller loggers used to measure speed directly decreased descent speeds by up to 0.5 m s⁻¹ when compared to smaller depth-only loggers. Cormorants in Greenland seem to combine two searching strategies, one requiring low speed to scan the water column or benthos, and one requiring high speed to pursue prey. These two strategies depend on the two main habitats of their prey: pelagic or demersal.     

The diving behaviour of brooding king penguins (<Emphasis Type="Italic">Aptenodytes patagonicus</Emphasis>) from the Falkland Islands: variation in dive profiles and synchronous underwater swimming provide new insights into their foraging strategies

The diving behaviour of king penguins (Aptenodytes patagonicus) was studied on the Falkland Islands, where a small population (ca. 300 fledglings year^–1) is located at the geographical limit of their breeding range. King penguins rearing newly hatched chicks were equipped with time-depth recorders before leaving for sea. In total, 20,175 dives >3 m were recorded from 12 birds during 15 foraging trips with a mean duration of 5.7±2.3 days. The majority of the trips was directed up to 500 km to the northeast of the breeding colony in slope waters of, and oceanic waters beyond, the Patagonian shelf. Mean time spent underwater accounted for 42±9% of the foraging trip. Mean dive depth achieved was 55±16 m; maximum dive depth recorded was 343 m. Mean dive duration was 159±25 s; maximum dive duration was 480 s. The mean vertical distance covered was 140±65 km trip^–1; and on average birds covered 25 km day^–1. Synchronous diving behaviour was observed in two birds for a period of about 24 h after leaving the colony. Dive depth correlated positively with: (1) light intensity, (2) dive duration and (3) vertical velocities, thus confirming previous findings obtained from conspecifics at other breeding sites and indicating comparable diving behaviour. However, separation of dives according to their profile—V-, U-, or W-shaped—revealed significant differences between certain dive parameters. For a given depth range, bottom time was longer and vertical velocities higher in W-dives than in U-dives. This, together with a higher number of W-dives at dawn and dusk, suggests that foraging is more effective during W-dives than U-dives, and during twilight. These findings imply that king penguins have to make more complex decisions, individually and socially, on the performance of the subsequent dive than previously thought.Communicated by O. Kinne, Oldendorf/Luhe     

Diving characteristics of southern rockhopper penguins (Eudyptes c. chrysocome) in the southwest Atlantic

The diving behaviour of southern rockhopper penguins (Eudyptes c. chrysocome) was studied at two breeding sites in the Southwest Atlantic: the Falkland Islands and Staten Island, Argentina. Incubating and brooding birds were equipped with time-depth recorders to monitor their foraging activities. Rockhopper penguins from Staten Island started their breeding season about 3 weeks earlier than their conspecifics from the Falkland Islands. The foraging area used by incubating males from the Falkland Islands comprised about 150,000 km² to the northeast of the breeding site and was characterised by shelf and slope waters, whereas the foraging area of incubating males from Staten Island comprised 350,000 km² of oceanic waters to the southeast of the breeding site. A number of dive parameters were measured and compared between the four study groups: Incubating males and brooding females from the Falkland Islands, and incubating males and females from Staten Island. In all study groups, dive depth correlated positively to light intensity, dive duration and vertical velocity. However, significant differences between various diving parameters of the study groups were noted, not only in terms of diving performance, but also as regards diving efficiency (DE). A principal component analysis (PCA) on 16 variables revealed that 75% of the variance could be explained by only two principal components: diving pattern (PC1) and diving effort (PC2). PC1 indicated that the birds from Staten Island, both males and females, dived deeper, covered a greater vertical distance per hour and had higher ascent rates, but spent less time underwater and at the bottom of a dive, and had a lower DE than conspecifics from the Falkland Islands. PC2, which included the percentage of foraging dives, the number of dives per hour, dive duration, bottom time and descent rate, differed significantly between incubating males from the Falkland Islands and the other three groups, which were all very similar. Overall, the diving behaviour was notably similar to that of conspecifics from the Indian and Pacific Oceans. The implications of the results in terms of intra-specific adaptations as well as potential threats from human activities are discussed.     

Depth utilisation by breeding Adélie penguins,Pygoscelis adeliae,at Esperanza Bay,Antarctica

Miniature depth gauges were attached in December 1987 and January 1988 to Adélie penguins,Pygoscelis adeliae, breeding at Esperanza on the Antarctic Peninsula. Results from 34 birds showed that foraging penguins with eggs and with brooded and crèching chicks spent mean periods away from the nest of 96, 36 and 21 h, respectively, during which time means of 29.0 h (30%), 11.2 h (31%) and 2.7 h (13%), respectively, were spent under water at depths > 5 m. Time under water was positively correlated with time absent from nest. Maximum depth reached was 170 m but overall birds spent most time at shallower depths. Birds foraging for brooded chicks dived deeper than birds foraging for crèching chicks. Stomach-pumping indicated that the principal prey caught at this time was krill,Euphausia superba. Mean mass changes of adults during single foraging trips indicated that krill were caught at a mean rate of 7.2 g min^–1 spent under water.     

Crossing the frontier: vertical transit rates of deep diving cormorants reveal depth zone of neutral buoyancy

Knowing the depth zone of neutral buoyancy of divers is important because buoyancy can determine how animals manage their energy budget. In this study, we estimate the depth zone of neutral buoyancy of free-ranging cormorants for the first time, using time-depth recorders. We discovered that vertical ascent rates of 12 Crozet and 15 Kerguelen diving blue-eyed shags (respectively Phalacrocorax melanogenis and P. verrucosus) slowed down considerably at the 50–60 m depth zone. We suggest this was due to birds trying to reach the surface from that point upwards using reduced locomotor activity because the force of buoyancy becomes greater than the force of gravity at that depth. The results show a shift of this depth zone in relation to maximum targeted dive depth, suggesting cormorants may control buoyancy through respiratory air volume adjustment. Interestingly, 60 m is close to the maximum depth zone reached by these two species during dives lasting 4 min, their estimated behavioural aerobic dive limit. This suggests that the decision to swim deeper has a direct consequence on the energy budget, with time spent recovering at the surface (time thus lost to foraging) strongly increasing relative to the preceding time of submergence. Resources found in deeper waters must be of sufficient quantity or quality to justify crossing the frontier of physical neutral buoyancy.     

Do activity costs determine foraging tactics for an arctic seabird?

How energy costs affect foraging decisions is poorly understood for marine animals. To provide data relevant to this topic, we examined the relationship between activity levels and foraging behavior by attaching activity recorders to 29 chick-rearing wing-propelled diving birds (thick-billed murres, Uria lomvia) in 1999–2000. We connected the activity during the final dive bout with the prey item we observed being fed to the chicks. After accounting for changes in activity level with depth, activity was highest during the final dive of a dive bout, reflecting maneuvring during prey capture. Pelagic prey items, especially invertebrates (amphipods), were associated with higher depth-corrected activity, leading to shorter dives for a given depth (presumably due to higher oxygen consumption rates) and, thus, shorter search times (lower bottom time for a given depth). Pelagic prey items were likely captured during active pursuit, with the birds actively seeking and pursuing schooling mid-water prey. In contrast, benthic prey involved low activity and extended search times, suggesting that the birds slowly glided along the bottom in search for prey hidden in the sediments or rocks. We concluded that activity levels are important in determining the foraging tactics of marine predators. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

Ontogeny of early diving and foraging behavior of northern fur seal (Callorhinus ursinus) pups from Bering Island,Russia

The ontogeny of diving and foraging behavior of northern fur seal pups from a stable population on Bering Island, Russia, was recorded with animal-borne instruments during their first few months at sea, a critical period during their first year at sea. Thirty-five pups were instrumented with satellite-linked time-depth recorders and stomach temperature pills. Diving occurred predominantly at night with deeper and longer dives as the pups matured. Mean dive depths were correlated with lunar illumination, whereas mean dive durations were also correlated with time of day and sex. Foraging success did not differ between sexes, and there was no relationship between meal size (as indicated by feeding event duration and minimum stomach temperature) and lunar illumination fraction or maximum foraging depth. Although most pups were able to successfully forage within 3 days of starting their migration, the number of feeding events recorded each day remained low (mean 1.6 events day^?1). There was no indication of an appreciable increase in meal size after the first 2 weeks of the migration despite an increase in dive frequency and depth. The results are consistent with observations that pups do not gain mass during their first year and emphasize the risk of starvation from infrequent foraging in cold water.     

Going deep: common murres dive into frigid water for aggregated,persistent and slow-moving capelin

Owing to the necessity of delivering food to offspring at colonies, breeding seabirds are highly constrained in their foraging options. To minimize constraints imposed by central-place foraging and to optimize foraging behavior, many species exhibit flexible foraging tactics. Here we document the behavioral flexibility of pursuit-diving common murres Uria aalge when foraging on female capelin Mallotus villosus in the northwest Atlantic. Quite unexpectedly, being visual foragers, we found that common murres dived throughout the day and night. Twenty-one percent of recorded dives (n = 272 of 1,308 dives) were deep (≥50 m; maximum depth = 152 m, maximum duration = 212 s), bringing murres into sub-0°C water in the Cold Intermediate Layer (CIL; 40–180 m) of the Labrador Current. Deep dives occurred almost exclusively during the day when murres would have encountered spatially predictable aggregations of capelin between 100 and 150 m in the water column. Temperatures within the CIL shaped trophic interactions and involved trade-offs for both predators and prey. Sub-0°C temperatures limit a fish’s ability to escape from endothermic predators by reducing burst/escape speeds and also lengthening the time needed to recover from burst-type activity. Thus, while deep diving may be energetically costly, it likely increases certainty of prey capture. Decreased murre foraging efficiency at night (indicated by an increase in the number of dives per bout) reflects both lower light conditions and changing prey behavior, as capelin migrate to warmer surface waters at night where their potential to escape from avian predators could increase.     

Chinstrap penguins alter foraging and diving behavior in response to the size of their principle prey, Antarctic krill

Penguins may exhibit plasticity in their diving and foraging behaviors in response to changes in prey availability. Chinstrap penguins are dependent predators of Antarctic krill in the Scotia Sea region, but krill populations have fluctuated in recent years. We examined the diet of chinstrap penguins at Livingston Island, South Shetland Islands, in relation to their diving and foraging behavior using time-depth recorders over six breeding seasons: 2002–2007. When krill were smaller, more chinstrap penguins consumed fish. In these years, chinstrap penguins often exhibited a shift to deep dives after sundown, and then resumed a shallower pattern at sunrise. These night dives were unexpectedly deep (up to 110 m) and mean night dive depths sometimes exceeded those from the daytime. The average size of krill in each year was negatively correlated to mean night dive depths and the proportion of foraging trips taken overnight. Based on these patterns, we suggest that when krill were small, penguins increasingly targeted myctophid fish. The average krill size was negatively correlated to the time chinstrap penguins spent foraging which suggests that foraging on smaller krill and fish incurred a cost: more time was spent at sea foraging.     

Prey ecology and behaviour affect foraging strategies in the Great Cormorant

The fine link between a particular dive pattern and a specific prey item represents a challenging task in the analysis of marine predator–prey relationships. There is growing evidence that prey type affects diving seabirds’ foraging strategies, dive shapes and underwater activity costs. This study investigates whether a generalist diver, the Great Cormorant Phalacrocorax carbo, modifies the time budget allocated to prey-capture behaviour and breathing strategies (reactive vs. anticipatory) with respect to the prey type (pelagic vs. benthic). Video recordings of 91 Great Cormorants show how the ecology and behaviour of their main prey, Mullets (Mugilidae) and Flounders Platichthys flesus, affect dive/surface durations and the diving pattern. The demersal habit and the low mobility of Flounders leads to an easy access to prey with an anticipatory strategy. Moreover, the patchy distribution of this fish species increases prey-capture rates. Conversely, Mullets exploit the whole water column and are highly mobile, and this is reflected in the need of performing two sequential dives to capture a prey, both longer and likely more expensive, with a consequent switch of strategy from reactive in the searching phase to anticipatory breathing during prey-capture events. This study provides evidence that a generalist diver may switch between different foraging strategies, and it shows how each of them may be optimal under particular ecological conditions. These constraints influence the dynamics that operate within the marine food chains and have relevant implications in managing lagoon areas, including fish ponds.     

Comparison of diving behavior and foraging habitat use between chinstrap and gentoo penguins breeding in the South Shetland Islands, Antarctica

Chinstrap, Pygoscelis antarctica, and gentoo, P. papua, penguins are sympatric species that inhabit the Antarctic Peninsula. To evaluate differences in the foraging habitat of these two species, we recorded their foraging locations and diving behavior using recently developed GPS-depth data loggers. The study was conducted on King George Island, Antarctica during the chick-guarding period of both species, from December 2006 to January 2007. The area used for foraging, estimated as the 95% kernel density of dive (>5 m) locations, overlapped partially between the two species (26.4 and 68.5% of the area overlapped for chinstrap and gentoo penguins, respectively). However, the core foraging area, estimated as the 50% kernel density, was mostly separate (12.8 and 25.0% of the area overlapped for chinstrap and gentoo penguins, respectively). Chinstrap penguins tended to use off-shelf (water depth > 200 m) regions (77% of the locations for dives >5 m), whereas gentoo penguins mainly used on-shelf (water depth < 200 m) areas (71% of dive locations). The data on foraging locations, diving behavior, and bathymetry indicated that gentoo penguins often performed benthic dives (28% of dives >5 m), whereas chinstrap penguins almost always used the epipelagic/mid-water layer (96% of dives >5 m). Diving parameters such as diving bottom duration or diving efficiency differed between the species, reflecting differences in the use of foraging habitat. The diving parameters also suggested that the on-shelf benthic layer was profitable foraging habitat for gentoo penguins. Conversely, the relationship between trip duration, date, and stomach content mass suggested that the chinstrap penguins went further from the colony to forage as the season progressed, possibly reflecting a reduction in prey availability near the colony. Our results suggest that chinstrap and gentoo penguins segregated their foraging habitat in the Antarctic coastal marine environment, possibly due to inter- and intra-specific competition for common prey resources.     

Archival tagging of subadult and adult common thresher sharks (Alopias vulpinus) off the coast of southern California

The common thresher shark (Alopias vulpinus) is a secondary target species of the California drift gillnet fishery (CA-DGN) and supports a growing recreational fishery in California waters. This study used archival tags to examine the movement patterns and habitat preferences of common threshers of the size range captured in the CA-DGN (>120 cm fork length). Depth and temperature-logging archival tags were deployed on 57 subadult and adult common threshers in the Southern California Bight. Tags from five individuals (8.8%) were recovered, and 154 days of data were successfully obtained from four of these. By night, shark movements were primarily limited to waters above the thermocline, which ranged in depth from 15 to 20 m. Sharks were significantly deeper by day, and daytime vertical distribution consisted of two distinct modes: a ‘shallow mode’ (wherein sharks occupied only the upper 20 m of the water column) and a ‘deep mode’ (characterized by frequent vertical excursions below the thermocline). This modal switch is interpreted as relating to regional differences in abundance of surface-oriented prey and prey in deeper water. Maximum dive depth was 320 m, greatest dive duration was 712 min, minimum temperature experienced during a dive was 9.1°C, and dive descent rate was significantly greater than ascent rate. Sharks inhabited waters corresponding to a sea surface temperature range of 16 to 21°C. The nocturnal depth distribution of common threshers has implications for management of drift gillnet deployment depths in the CA-DGN.     

Lip-reading in remote subjects: an attempt to quantify and separate ingestion, breathing and vocalisation in free-living animals using penguins as a model

A new mandibular sensor is presented here based on the use of a Hall sensor, attached to one mandible, opposite a magnet, attached to the other mandible. Changes in sensor voltage, proportional to magnetic field strength, and thus inter-mandibular angle, are recorded in a logger. This system was tested on seven captive Adélie penguins (Pygoscelis adeliae) and three gentoo penguins (Pygoscelis papua) during: (1) feeding trials on land, where birds were given known quantities and types of food; and (2) trials in water where birds were allowed to swim and dive freely. In addition, six free-living Magellanic penguins (Spheniscus magellanicus) were equipped with the system for single foraging trips. Angular signatures were looked for in instances when both captive and free-living birds might open their beaks, and it was discovered that five major behaviours could be identified: ingestion, breathing, calling, head shaking and preening. Captive feeding trials showed that prey mass could be determined with reasonable accuracy (r²=0.92), and there was some indication that prey type could be resolved if recording frequency were high enough. Vocalisations in Adélie penguins (arc calls) took <0.7 s for mean maximum beak angles of 4.2° (SD 1.3), and were distinguished by their relatively gradual change in beak angle and by their high degree of symmetry. Beak shakings were distinguishable by their short duration (multiple peaks of <0.5 s) and minimal maximum angle (<0.5°). Preening behaviour was apparent due to multiple decreasing peaks (angles <8°). Breathing could be subdivided into that during porpoising, where a characteristic double peak in beak angle was recorded, and that during normal surface rests between dives. During porpoising, only the primary peak (mean maximum beak angle 25.1°, SD 4.7) occurred when the bird was out of the water (mean maximum for second peak 5.9°, SD 4.1). During normal surface rests in free-living birds, breaths could be distinguished as a series of beak openings and closures, showing variation in amplitude and frequency according to an apparent recovery from the previous dive and preparation for the subsequent dive to come. The mandibular measuring system presented shows considerable promise for elucidating many hitherto intractable aspects of the behaviour of free-living animals.     

Water column use and forage strategies of female southern elephant seals from Marion Island

The at-sea behaviour of marine top predators provides valuable insights into the distribution of prey species and strategies used by predators to exploit patchily distributed resources. We describe the water column usage and dive strategies of female southern elephant seals from Marion Island tracked between 2004 and 2008. Dives representing increases in forage effort were identified using a method that combines dive type analyses and the calculation of relative amounts of time that animals spend in the bottom phases of dives. Results from this analysis indicate that female elephant seals from Marion Island tend to display lower levels of forage effort closer to the island and display intensive opportunistic forage bouts that occur at a minimum distance of approximately 215 km from the island. Females from Marion Island dived deeper and for longer periods of time, compared to females from other populations. Most animals displayed positive diel vertical migration, evidently foraging pelagically on vertically migrating prey. A few animals displayed periods of reverse (negative) diel vertical migration, however, diving to deeper depths at night, compared to daytime. This behaviour is difficult to explain and prey species targeted during such periods unknown. Our results illustrate plasticity in foraging behaviour of southern elephant seals, as well as inter-population differences in forage strategies.     

Contrasting foraging tactics by northern gannets (<Emphasis Type="Italic">Sula bassana</Emphasis>) breeding in different oceanographic domains with different prey fields

In order to forage and to provision offspring effectively, seabirds negotiate a complex of behavioural, energetic, environmental and social constraints. In first tests of GPS loggers with seabirds in North America, we investigated the foraging tactics of free-ranging northern gannets (Sula bassana) at a large and a medium-sized colony that differed in oceanography, coastal position and prey fields. Gannets at Low Arctic colony (Funk Island) 50 km off the northeast coast of Newfoundland, Canada provisioned chicks almost entirely with small forage fish (capelin Mallotus villosus, 89%), while at boreal colony (Bonaventure Island) 3 km from shore in the Gulf of St. Lawrence, Quebec, Canada, large pelagic fish dominated parental prey loads (Atlantic mackerel Scomber scombrus 50%, Atlantic herring Clupea harengus 33%). Mean foraging range and the total distance travelled per foraging trip were significantly greater at the larger inshore colony (Bonaventure) than at the smaller offshore colony (Funk Island; 138 and 452 km vs. 64 and 196 km, respectively). Gannets from Funk Island consistently travelled inshore to forage on reproductive capelin shoals near the coast, whereas foraging flights of birds from Bonaventure were much more variable in direction and destination. Birds from the Low Arctic colony foraged in colder sea surface water than did birds from the boreal colony, and dive characteristics differed between colonies, which is concordent with the difference in prey base. Differences between the colonies reflect oceanographic and colony-size influences on prey fields that shape individual foraging tactics and in turn generate higher level colony-specific foraging “strategies”.     

The influence of subsurface thermal structure on the diving behavior of northern fur seals (<Emphasis Type="Italic">Callorhinus ursinus</Emphasis>) during the breeding season

In the heterogeneous marine environment, predators can increase foraging success by targeting physical oceanographic features, which often aggregate prey. For northern fur seals (Callorhinus ursinus), two prevalent oceanographic features characterize foraging areas during summer in the Bering Sea: a stable thermocline and a subsurface “cold pool”. The objective of this study was to examine the influence of these features on foraging behavior by equipping fur seals from St. Paul Island (Alaska, USA) with time-depth recorders that also measured water temperature. Foraging bout variables (e.g., mean dive depth and percent time diving in a bout) were compared with respect to subsurface thermal characteristics (thermocline presence and strength and cold pool presence). Over 74% of bouts occurred in association with strong thermoclines (temperature change > 5°C). Few differences were found for dive behavior in relation to the presence of a thermocline and the cold pool, but for epipelagic bouts, a strong thermocline resulted in increased bottom times, number of dive wiggles, and percent time diving when compared to moderate thermoclines. There was also a positive relationship between mean dive depth and thermocline depth. The combination of increasing foraging effort in areas with strong thermoclines and diving to depths closely related to the thermocline indicates this feature is important foraging habitat for northern fur seals and may act to concentrate prey and increase foraging success. By recognizing the environmental features northern fur seals use to find prey, managers will be better equipped to identify and protect foraging habitat that is important to northern fur seals, and possibly other marine predators in the Bering Sea.