Linking marine predator diving behavior to local prey fields in contrasting habitats in a subarctic glacial fjord

Foraging theory predicts that animals will adjust their foraging behavior in order to maximize net energy intake and that trade-offs may exist that can influence their behavior. Although substantial advances have been made with respect to the foraging ecology of large marine predators, there is still a limited understanding of how predators respond to temporal and spatial variability in prey resources, primarily due to a lack of empirical studies that quantify foraging and diving behavior concurrently with characteristics of prey fields. Such information is important because changes in prey availability can influence the foraging success and ultimately fitness of marine predators. We assessed the diving behavior of juvenile female harbor seals (Phoca vitulina richardii) and prey fields near glacial ice and terrestrial haulout sites in Glacier Bay (58°40′N, ?136°05′W), Alaska. Harbor seals captured at glacial ice sites dived deeper, had longer dive durations, lower percent bottom time, and generally traveled further to forage. The increased diving effort for seals from the glacial ice site corresponded to lower prey densities and prey at deeper depths at the glacial ice site. In contrast, seals captured at terrestrial sites dived shallower, had shorter dive durations, higher percent bottom time, and traveled shorter distances to access foraging areas with much higher prey densities at shallower depths. The increased diving effort for seals from glacial ice sites suggests that the lower relative availability of prey may be offset by other factors, such as the stability of the glacial ice as a resting platform and as a refuge from predation. We provide evidence of differences in prey accessibility for seals associated with glacial ice and terrestrial habitats and suggest that seals may balance trade-offs between the costs and benefits of using these habitats.     

Geographical variation in the behaviour of a central place forager: Antarctic fur seals foraging in contrasting environments

Foragers show adaptive responses to changes within their environment, and such behavioural plasticity can be a significant driving force in speciation. We investigated how lactating Antarctic fur seals, Arctocephalus gazella, adapt their foraging within two contrasting ecosystems. Location and diving data were collected concurrently, between December 2003 and February 2004, from 43 seals at Bird Island, where krill, Euphausia superba, are the main prey, and 39 at Heard Island, where mostly fish are consumed. Seals at Heard Island were shorter and lighter than those at Bird Island and they spent longer at sea, dived more frequently and spent more time in the bottom phase of dives. Generalized additive mixed effects models showed that diving behaviours differed between the islands. Both populations exploited diel vertically migrating prey species but, on average, Heard Island seals dived deeper and exceeded their estimated aerobic dive limits. We propose that the recovery of the Heard Island population may be limited by the relative inaccessibility and scarcity of food, whereas at Bird Island, the presence of abundant krill resources helps sustain extremely high numbers of seals, even with increased intra- and inter-specific competition. Both populations of fur seals appear to be constrained by their physiological limits, in terms of their optimal diving behaviour. However, there does appear to be some flexibility in strategy at the level of trip with animals adjusting their time at sea and foraging effort, in order to maximize the rate of delivery of energy to their pups.     

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.     

Temporal changes in activity budgets of chick-rearing Adélie penguins

Changes in sea-ice conditions can affect locomotion on land, diving behavior, and corresponding foraging success of penguins. In this study, locomotion on land and diving behavior were compared between early and late stages of the guard phase with different sea-ice conditions using miniaturized time-depth-acceleration data loggers for Adélie penguins Pygoscelis adeliae from 18 December 2001 to 11 January 2002 in Dumont d’Urville, Adélie Land (66.7°S, 140.0°E), Antarctica. Differences were found between early and late stages in the ratio of walking vs. tobogganing, proportion of time spent diving, diving depth as well as in the rate of parental tissue accumulation. In contrast, trip duration, distance traveled on land, and meal delivery rate to chicks did not differ between the stages. This study suggests that physical changes in sea-ice during the penguins’ chick-rearing period may affect certain on-land and/or at-sea behaviors which, in turn, may affect how resources are allocated to self-maintenance or chick-provisioning.     

Evidence of a maternal foraging cycle resembling that of otariid seals in a small phocid,the harbor seal

Summary Lactation strategies in the two largest families of seals have been characterized as a phylogenetic dichotomy, with sea lions and fur seals (Otariidae) exhibiting foraging cycles and true seals (Phocidae) a strategy of fasting. We show that a lactating phocid, the harbor seal, Phoca vitulina, has a foraging cycle similar to that of otariids. Time-depth recorders attached to lactating harbor seal mothers revealed that 9 of 11 females began bouts of diving, averaging 12–40 m, by mid-lactation (12 days). During the remainder of lactation, females made an average of seven diving trips, lasting about 7 h. They returned to the rookery during the interval between successive bouts to nurse their pups. Diving was more frequent during daylight than at night and diving bouts increased in duration as lactation progressed. The diving behavior of females that had weaned their pups and previously collected data from stomach lavage, suggest that the bouts of diving represent successful foraging. We propose that the lactation strategy of the harbor seal is intermediate to that of the otariids and other phocids studied. The harbor seal has a foraging cycle like the otariids, but typically resembles other phocids in length of lactation, rate of mass gain in pups, and in milk fat content. As harbor seals are among the smallest phocids, and only slightly larger than most otariids, it seems likely that maternal size constrains the amount of stored energy harbor seal females can bring to the rookery, forcing them to start feeding during the lactation period.Correspondence to: D.J. Boness     

Foraging behaviour of sympatric Antarctic and subantarctic fur seals: does their contrasting duration of lactation make a difference?

The duration of periods spent ashore versus foraging at sea, diving behaviour, and diet of lactating female Antarctic (Arctocephalus gazella, AFS) and subantarctic (A. tropicalis, SFS) fur seals were compared at Iles Crozet, where both species coexist. The large disparity in lactation duration (SFS: 10 months, AFS: 4 months), even under local sympatry, has led to the expectation that AFS should exhibit higher foraging effort or efficiency per unit time than SFS to allow them to wean their pups in a shorter period of time. Previous evidence, however, has not supported these expectations. In this study, the distribution of foraging trip durations revealed two types of trips: overnight (OFT, <1 day) and long (LFT, >1 day), in common with other results from Macquarie Island. However, diving behaviour differed significantly between foraging trip types, with greater diving effort in OFTs than in LFTs, and diving behaviour differed between fur seal species. OFTs were more frequent in SFS (48%) than in AFS (28%). SFS performed longer LFTs and maternal attendances than AFS, but spent a smaller proportion of their foraging cycle at sea (66.2 vs. 77.5%, respectively). SFS dove deeper and for longer periods than AFS, in both OFTs and LFTs, although indices of diving effort were similar between species. Diel variation in diving behaviour was lower among SFS, which foraged at greater depths during most of the night time available than AFS. The diving behaviour of AFS suggests they followed the nychthemeral migration of their prey more closely. Concomitant with the differences in diving behaviour, AFS and SFS fed on the same prey species, but in different proportions of three myctophid fish (Gymnoscopelus fraseri, G. piabilis, and G. nicholsi) that represented most of their diet. The estimated size of the most important fish consumed did not vary significantly between fur seal species, suggesting that the difference in dive depth was mostly a result of changes in the relative abundance of these myctophids. The energy content of these fish at Iles Crozet may thus influence the amount and quality of milk delivered to pups of each fur seal species. These results contrast with those found at other sites where both species coexist, and revealed a scale of variation in foraging behaviour which did not affect their effort while at sea, but that may be a major determinant of foraging efficiency and, consequently, maternal investment.     

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.     

Variation in foraging effort by lactating Antarctic fur seals: response to simulated increased foraging costs

Seasonally breeding predators, which are limited in the time available for provisioning young at a central location, and by the fasting abilities of the young, are likely to maximize energy delivery to the young by maximizing the rate of energy delivery averaged over the whole period of investment. Reduction in food availability or increased foraging costs will alter the optimal behavior of individuals. This study examined the behavioral adaptations of a diving predator, the Antarctic fur seal, to increased foraging costs during lactation. One group of mothers (n=5, treatment) was fitted with additional drag to increase the cost of transport in comparison with a control group (n=8). At the scales of the individual dives, the treatment group made more shorter, shallower (< 30 m) dives. Compensation for slower swimming speeds was achieved by diving at a steeper angle. Overall, diving behavior conformed to several specific theoretical predictions but there were also departures from theory, particularly concerning swimming speed during diving. Diving behavior appears to be adjusted to maximize the proportion of time spent at the bottom of dives. At the scale of diving bouts, no difference was observed between the treatment and control groups in terms of the frequency and duration of bouts and there was also no difference between the two groups in terms of the proportion of time spent diving. At the scale of complete foraging cycles, time taken to return to the pup was significantly longer in the treatment group but there was no difference in the rate of delivery of energy (measured from pup growth rate) to the pups in each group. Since mothers in the treatment group did not use significantly more body reserves, we conclude that behavioral adjustments at the scale of individual dives allowed mothers in the treatment group to compensate for the additional foraging costs. Pup growth rate appears to be less sensitive to the foraging conditions experienced by mothers than foraging trip duration. Received: 14 June 1996 / Accepted after revision: 16 November 1996     

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     

Energy expenditure and food consumption of foraging Imperial cormorants in Patagonia, Argentina

Energy management during the breeding season is crucial for central place foragers since parents need to feed themselves and their offspring while being spatially and temporally constrained. In this work, we used overall dynamic body acceleration as a measure of activity and also to allude to the foraging energy expenditure of breeding Imperial cormorants Phalacrocorax atriceps. We also analyzed how changes in the time or energy allocated to different activities affected the foraging trip energy expenditure and estimated the daily food requirements of the species. Birds spent 42 % of the total energy flying to and from the feeding areas and 16 % floating at sea. The level of activity underwater was almost 1.5 times higher for females than for males. The most expensive diving phase in terms of rate of energy expenditure was descending though the water column. The total foraging trip energy expenditure was particularly sensitive to variation in the amount of time spent flying. During the breeding season, adult cormorants breeding along the Patagonian coast would consume approximately 10,000 tons of food.     

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.     

In situ measures of foraging success and prey encounter reveal marine habitat-dependent search strategies

Predators are thought to reduce travel speed and increase turning rate in areas where resources are relatively more abundant, a behavior termed "area-restricted search." However, evidence for this is rare, and few empirical data exist for large predators. Animals exhibiting foraging site fidelity could also be spatially aware of suitable feeding areas based on prior experience; changes in movement patterns might therefore arise from the anticipation of higher prey density. We tested the hypothesis that regions of area-restricted search were associated with a higher number of daily speed spikes (a proxy for potential prey encounter rate) and foraging success in southern elephant seals (Mirounga leonina), a species exhibiting both area-restricted searches and high interannual foraging site fidelity. We used onshore morphological measurements and diving data from archival tags deployed during winter foraging trips. Foraging success was inferred from in situ changes in relative lipid content derived from measured changes in buoyancy, and first-passage time analysis was used to identify area-restricted search behavior. Seals exhibited relatively direct southerly movement on average, with intensive search behavior predominantly located at the distal end of tracks. The probability of being in search mode was positively related to changes in relative lipid content; thus, intensively searched areas were associated with the highest foraging success. However, there was high foraging success during the outward transit even though seals moved through quickly without slowing down and increasing turning rate to exploit these areas. In addition, the probability of being in search mode was negatively related to the number of daily speed spikes. These results suggest that movement patterns represent a response to prior expectation of the location of predictable and profitable resources. Shelf habitat was 4-9 times more profitable than the other habitats, emphasizing the importance of the East Antarctic shelf for this and other predators in the region. We have provided rare empirical data with which to investigate the relationship between predator foraging strategy and prey encounter/ foraging success, underlining the importance of inferring the timing and spatial arrangement of successful food acquisition for interpreting foraging strategies correctly.     

Winter distribution and haul-out behaviour of female Antarctic fur seals from South Georgia

Telemetry-based techniques have revealed the foraging patterns of many land breeding marine predators, especially during the summer breeding season. However, during the winter, when freed from the constraints of provisioning their young, such animals are more difficult to track. Using geolocation (Global Location Sensing, GLS) loggers and satellite tags (Platform Terminal Transmitters, PTTs) we successfully tracked 16 female Antarctic fur seals from South Georgia during the austral winter. The majority of females concentrated their winter foraging in the waters around the breeding beaches (90% of locations were within 510 km). However, as the winter progressed, two of the seals spent a number of months to the south, in and around the seasonal ice edge, and five seals migrated north and northwest from South Georgia. Four of these seals clearly crossed the Polar Front and two reached the Patagonian Shelf, apparently exploiting the continental shelf edge and the Subantarctic Front. Activity (saltwater immersion) data suggested that seals spent the majority of the winter months at sea but there were rare occasions when seals hauled out, either on land or on ice floes. We obtained data from two individuals that enabled us to compare the performance of PTT and GLS devices. For these seals the mean distance between GLS and PTT locations was 122 and 132 km. Although the recovery rates were low in this study, given improvements in attachment techniques, we have demonstrated that these micro-geolocation loggers provide an ideal tool with which to study the long-term dispersal of diving marine predators at larger scales. This is the first study to show that female fur seals from South Georgia remain at sea for almost the entirety of the non-breeding winter period. Using land-based observations it has been assumed that the fur seal population at South Georgia has little temporal overlap with the krill fishery that operates mostly during the winter months in this region. We have shown that a large proportion of the female fur seals that breed on South Georgia potentially remain in the vicinity of the island and are thus in direct competition with the region’s fisheries activities.     

Size and seasonal influences on the foraging range of female grey seals in the northeast Atlantic

Evidence of segregation in foraging habitat has been demonstrated in some top marine predators, including cetaceans, pinnipeds and seabirds. However, most data are not adequate to assess differences relating to body size or seasonal influences. This has implications for quantitative modelling of population-level predator–prey interactions and ecosystem structure. We examined potential influence of body size and ‘fatness’ on the foraging trip characteristics of a top marine predator, the Atlantic grey seal (Halichoerus grypus), in southwest Ireland within the framework of optimal foraging theory to examine how female grey seals foraging behaviour varied with size, and across the period between moult and breeding. Larger seals undertook trips of greater duration and travelled further from haul-out sites than smaller seals. However, body fat was negatively associated with trip duration and extent. Seals spent more time at sea during the summer, but trips were shorter in extent, suggesting more localized foraging during this season.     

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.     

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.     

Diving deep in a foraging hotspot: acoustic insights into bottlenose dolphin dive depths and feeding behaviour

To exploit resources in their environment, odontocete cetaceans have evolved sophisticated diving abilities to allow effective foraging. However, data on the diving behaviour and underwater foraging behaviour remains limited. This study made use of echolocation clicks and other calls to study the diving behaviour of bottlenose dolphins. Dolphins used the full water column and consistently dived to depths of around 50 m, close to the seabed. However, the majority of their time appeared to be spent within the surface layers of the water column. In addition, by localising calls that have been associated with prey capture events (Janik, Proc R Soc Lond Ser B 267:923–927, 2000a), it appeared that certain forms of feeding behaviour occurred primarily at depths of between 20 and 30 m. Furthermore, data on the depth of clicks made before and after these feeding calls suggested that during the minute before the calls, dolphins were consistently diving from the surface to depths close to the seabed, and were subsequently returning to the surface after the calls. This passive acoustic technique proved an accurate method for studying the depth distribution of dolphin vocalisations. By exploiting the natural sounds made by these wild odontocetes, this investigation provided a previously unavailable perspective on the the 3D nature of bottlenose dolphins foraging behaviour. It confirmed that while the dolphins spent the majority of time close to the surface, the full water column was exploited during foraging events.     

Spatial position and feeding success in ring-tailed coatis

The location of an animal within a social group has important effects on feeding success. When animals consume quickly eaten food items, individuals located at the front edge of a group typically have greater foraging success. When groups feed at large clumped resources, dominant individuals can often monopolize the resource, leading to higher feeding success in the center of the group. In order to test these predictions, behavioral data relating foraging success to within-group spatial position were recorded from two habituated groups of ring-tailed coatis (Nasua nasua) in Iguazu, Argentina. Foraging success did not fit expected patterns. When feeding on small ground litter invertebrates, coatis had the same foraging success at all spatial positions. This pattern likely resulted from an abundance of invertebrates in the ground litter. When feeding on fruit, individuals in the front of the group had greater feeding success, which was driven by the relatively quick depletion of fruit trees. Dominant juveniles were often located in the front of the group which led to increased access to food. This resulted in higher feeding success on fruits but simultaneously increased their risk of predation. Although groups typically became more elongated and traveled faster when feeding on fruit, it did not appear that the coatis were drastically changing their spacing strategies when switching between the two food types. Paradoxically, spatial position preferences during invertebrate foraging appeared to be driven by fruit trees. Because fruit trees were encountered so frequently, juveniles ranging at the front edge of the group during invertebrate foraging were the first to arrive at fruit trees and thus had higher foraging success. This study demonstrates the importance of how food patch size and depletion rate affect the spatial preferences of individuals.     

Factors affecting the foraging behaviour of the European shag: implications for seabird tracking studies

Seabird tracking has become an ever more popular tool to aid environmental procedures such as the designation of marine protected areas and environmental impact assessments. However, samples used are usually small and little consideration is given to experimental design and sampling protocol. European shags Phalacrocorax aristotelis were tracked using GPS technology over three breeding seasons and the following foraging trip characteristics: trip duration, trip distance, maximum distance travelled from the colony, size of area used and direction travelled from colony were determined for each foraging trip. The effect of sex, year of study, breeding site, number and age of chicks and the timing of tracking on foraging behaviour were investigated using a General Estimation Equation model. A range of sampling scenarios reflecting likely field sampling were also tested to compare how foraging behaviour differed depending on composition of the sample of birds tracked. Trip distance, trip duration, maximum distance travelled and size of area used were all significantly affected by the breeding site, and the number of chicks a tracked adult was raising. The effect of sex was also seen when examining trip distance, trip duration and the maximum distance travelled. The direction travelled on a foraging trip was also significantly affected by breeding site. This study highlights the importance of sampling regime and the influence that year, sex, age, number of chicks and breeding site can have on the foraging trip characteristics for this coastal feeding seabird. Given the logistical and financial constraints in tracking large numbers of individuals, this study identifies the need for researchers to consider the composition of their study sample to ensure any identified foraging areas are as representative as possible of the whole colony’s foraging area.     

An energy–distance trade-off in a central-place forager,the Antarctic fur seal (<Emphasis Type="Italic">Arctocephalus gazella</Emphasis>)

We tested the prediction that lactating fur seals (Arctocephalus gazella) at South Georgia will take prey of greater energy density with increasing distance of foraging from the colony. The study investigated the differences in diet of fur seals foraging within two regions, one near the breeding colony and the other at greater distance. Diet varied significantly in relation to foraging location. Dietary items of low quality were eaten in both regions but more food items with a high-energy content appeared in the diet of seals travelling to distant oceanic waters. We conclude that there is likely to be a trade-off between energy gain and distance travelled which enables female fur seals to maintain a relatively constant rate of energy delivery to their offspring irrespective of the distance travelled to find food.