A Bayesian hierarchical model of Antarctic fur seal foraging and pup growth related to sea ice and prey abundance

We created a Bayesian hierarchical model (BHM) to investigate ecosystem relationships between the physical ecosystem (sea ice extent), a prey measure (krill density), predator behaviors (diving and foraging effort of female Antarctic fur seals, Arctocephalus gazella, with pups) and predator characteristics (mass of maternal fur seals and pups). We collected data on Antarctic fur seals from 1987/1988 to 1994/1995 at Seal Island, Antarctica. The BHM allowed us to link together predators and prey into a model that uses all the data efficiently and accounts for major sources of uncertainty. Based on the literature, we made hypotheses about the relationships in the model, which we compared with the model outcome after fitting the BHM. For each BHM parameter, we calculated the mean of the posterior density and the 95% credible interval. Our model confirmed others' findings that increased sea ice was related to increased krill density. Higher krill density led to reduced dive intensity of maternal fur seals, as measured by dive depth and duration, and to less time spent foraging by maternal fur seals. Heavier maternal fur seals and lower maternal foraging effort resulted in heavier pups at 22 d. No relationship was found between krill density and maternal mass, or between maternal mass and foraging effort on pup growth rates between 22 and 85 days of age. Maternal mass may have reflected environmental conditions prior to the pup provisioning season, rather than summer prey densities. Maternal mass and foraging effort were not related to pup growth rates between 22 and 85 d, possibly indicating that food was not limiting, food sources other than krill were being used, or differences occurred before pups reached age 22 d.     

Are penguins and seals in competition for Antarctic krill at South Georgia?

The Antarctic fur seal (Arctocephalus gazella) and macaroni penguin (Eudyptes chrysolophus) are sympatric top predators that occur in the Southern Ocean around South Georgia where they are, respectively, the main mammal and bird consumers of Antarctic krill (Euphausia superba). In recent years the population of fur seals has increased, whereas that of macaroni penguins has declined. Both species feed on krill of similar size ranges, dive to similar depths and are restricted in their foraging range at least while provisioning their offspring. In this study we test the hypothesis that the increased fur seal population at South Georgia may have resulted in greater competition for the prey of macaroni penguins, leading to the decline in their population. We used: (1) satellite-tracking data to investigate the spatial separation of the Bird Island populations of these two species whilst at sea during the breeding seasons of 1999 and 2000 and (2) diet data to assess potential changes in their trophic niches between 1989 and 2000. Foraging ranges of the two species showed considerable overlap in both years, but the concentrations of foraging activity were significantly segregated spatially. The size of krill taken by both species was very similar, but over the last 12 years the prevalence of krill in their diets has diverged, with nowadays less krill in the diet of macaroni penguins than in that of Antarctic fur seals. Despite a significant degree of segregation in spatial resource use by the study populations, it is likely that the South Georgia populations of Antarctic fur seal and macaroni penguin exploit the same krill population during their breeding season. For explaining the opposing population trends of the two species, the relative contributions of independent differential response to interannual variation in krill availability and of interspecies competition cannot be resolved with available evidence. The likely competitive advantage of Antarctic fur seals will be enhanced as their population continues to increase, particularly in years of krill scarcity.     

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.     

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.     

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.     

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.     

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.     

Diving behaviour of Little Penguins from four colonies across their whole distribution range: bathymetry affecting diving effort and fledging success

Little Penguins, Eudyptula minor, breed in several small colonies in New Zealand and Australia. In this study, we compare the birds’ diving performances at different sites situated throughout their breeding range. Environmental conditions and breeding success vary drastically amongst colonies, but all birds feed on similar types of prey and face similar limitations on their foraging range. We examined several diving parameters and calculated the proportion of foraging zone available during breeding to examine whether oceanographic and geographic factors in the foraging zone can explain variations in diving behaviour and fledging success among the different colonies. In colonies with high fledging success, Penguin Island and Oamaru, penguins made shallow dives <50 m depth and had lower diving effort. More than 90% of the foraging zone was in waters <50 m depth in these colonies. Motuara Island also has shallow waters with 95% <50 m depth, but the fledging success was low. Phillip Island has only 42% of waters <50 m and comparatively low fledging success. Thus, penguins dived deeper and showed a higher diving effort in colonies with lower fledging success (Motuara Island and Phillip Island), indicating that they were disadvantaged compared to conspecifics from other colonies that dived shallower and with a lesser diving effort. We concluded that bathymetry is an important factor, but not the only one, which influences fledging success.     

Maternal investment in Antarctic fur seals: evidence for equality in the sexes?

Studies of the otariids (fur seals and sea lions), a highly sexually dimorphic group, have provided conflicting evidence of differential maternal expenditure in male and female offspring and, thus, suggestions that they conform to predictions of investment theory are equivocal. Since the mid-1970s, a diversity of research on Antarctic fur seals (Arctocephalus gazella) including studies of their reproductive ecology, lactation energetics, and foraging behaviour have been conducted at Bird Island, South Georgia that have resulted in one of the more complete and diverse data sets for any species of otariid. These long-term data were reviewed to determine whether there was any evidence to support that differential maternal expenditure occurred in Antarctic fur seals. Most of the data examined were collected during five consecutive austral summers from 1988 through 1992 and included years in which local food resources were abundant and scarce. We were unable to detect differences in the sex ratios of pups at birth or sex-biased differences in growth rates estimated from serial data, the number of foraging trips made, the duration of attendance ashore, diving behaviour, suckling behaviour, or milk consumption in any year and in the duration of foraging trips or age at weaning in 2 of 3 years. In addition, we found no evidence of greater reproductive costs between mothers with sons or daughters relative to their reproductive performance the following year. In contrast, sex-biased differences were only found in the duration of foraging trips in 1990, the age at weaning in 1988, and consistently in growth rates estimated from cross-sectional data. We suggest that differential maternal expenditure does not occur in Antarctic fur seals because male pups probably do not gain greater benefit from additional maternal expenditure than female pups. After weaning, males experience a period of rapid juvenile growth over 3–4 years during which time body mass nearly trebles. This growth will almost certainly be dependent upon available food resources then rather than on any maternal expenditure received over the first 4 months of life and, thus, the assumptions of the Trivers and Willard hypothesis are probably invalid for Antarctic fur seals. Received: 10 July 1996 / Accepted after revision: 3 March 1997     

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.     

Can marine mammals be used to monitor oceanographic conditions?

The breeding performance of higher predators has often been used to monitor fluctuations in the abundance of important prey stocks in marine ecosystems. The development of electronic data-loggers in recent years has also provided the opportunity of using wide-ranging marine animals to measure physical oceanographic conditions. In this study, time–depth recorders (TDRs) programmed to record temperature were deployed on female Antarctic fur seals (Arctocephalus gazella) at Bird Island, South Georgia (54°00′S; 38°02′W) during the breeding seasons 1994 to 1998. Temperature sensors had relatively slow response times, and thermal radiation errors occurred during the day when seals spent a large proportion of their time at the surface. Nevertheless, measurements provided temperature–depth profiles which were typical of the vertical stratification of the ocean. During the early stages of a foraging trip temperature increased, suggesting that fur seals travelled northwards from South Georgia towards the warmer waters of the Polar Front. In addition, higher temperatures were recorded by females that remained at sea for longer, implying that these individuals also travelled further. Mean sea-surface temperature (SST) increased from ∼1 to 4 °C from December to March and agreed with SSTs from ship, buoy and satellite. Future studies on marine mammals which combine satellite tracking with oceanographic measurements are likely to provide valuable information on biophysical aspects of the ocean. Received: 16 June 1998 / Accepted: 13 February 1999     

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.     

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     

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.     

Flexible foraging strategies of gentoo penguins <Emphasis Type="Italic">Pygoscelis papua</Emphasis> over 5 years in the South Shetland Islands,Antarctica

Gentoo penguins Pygoscelis papua show considerable plasticity in their diet, diving, and foraging behaviors among colonies; we expected that they might exhibit similar variability over time, at a single site, since flexible foraging habits would provide a buffer against changes in prey availability. We examined interannual changes in the foraging strategies and diet of gentoo penguins in the South Shetland Islands, Antarctica, over 5 years with variable prey abundance. Antarctic krill Euphausia superba was the primary diet item, and fish the secondary, though the importance of these items varied among years. Diving behavior also varied over time: different dive depth distributions were observed among years. Nonetheless, chick-rearing success remained relatively constant, indicating that gentoo penguins were able to maintain chick provisioning by altering their foraging strategy among years. Variable abundance of krill in the region did not have observable impacts on the diet, foraging behaviors or chick-rearing success of gentoo penguins. We suggest that foraging plasticity may be one reason that gentoo penguin populations have remained stable in the region, while their congeners (P. antarctica and P. adeliae) with less flexible foraging strategies have declined.     

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.     

Linking predator and prey behaviour: contrasts between Antarctic fur seals and macaroni penguins at South Georgia

Antarctic fur seals Arctocephalus gazella and macaroni penguins Eudyptes chrysolophus are the two main land-based krill Euphausia superba consumers in the northern Scotia Sea. Using a combination of concurrent at-sea (predator observations, net hauls and multi-frequency acoustics), and land-based (animal tracking and diet analysis) techniques, we examined variability in the foraging ecology of these sympatric top predators during the austral summer and autumn of 2004. Krill availability derived from acoustic surveys was low during summer, increasing in autumn. During the breeding season, krill occurred in 80% of fur seal diet samples, with fish remains in 37% of samples. Penguin diets contained the highest proportion of fish in over 20 years of routine monitoring (46% by mass; particularly the myctophid Electrona antarctica), with krill (33%) and amphipods (Themisto gaudichaudii; 21%) also occurring. When constrained by the need to return and feed their offspring both predator species foraged to the northwest of South Georgia, consistent with an area of high macrozooplankton biomass, but fur seals were apparently more successful at exploiting krill. When unconstrained by chick-rearing (during March) penguins foraged close to the Shag Rocks shelf-break, probably exploiting the high daytime biomass of fish in this area. Penguins and seals are able to respond differently to periods of reduced krill abundance (in terms of variability in diet and foraging behaviour), without detriment to the breeding success of either species. This highlights the importance of myctophid fish as an alternative trophic pathway for land-based predators in the Scotia Sea ecosystem.     

Modelling predation by transient leopard seals for an ecosystem-based management of Southern Ocean fisheries

Correctly quantifying the impacts of rare apex marine predators is essential to ecosystem-based approaches to fisheries management, where harvesting must be sustainable for targeted species and their dependent predators. This requires modelling the uncertainty in such processes as predator life history, seasonal abundance and movement, size-based predation, energetic requirements, and prey vulnerability. We combined these uncertainties to evaluate the predatory impact of transient leopard seals on a community of mesopredators (seals and penguins) and their prey at South Georgia, and assess the implications for an ecosystem-based management. The mesopredators are highly dependent on Antarctic krill and icefish, which are targeted by regional fisheries. We used a state-space formulation to combine (1) a mark-recapture open-population model and individual identification data to assess seasonally variable leopard seal arrival and departure dates, numbers, and residency times; (2) a size-based bioenergetic model; and (3) a size-based prey choice model from a diet analysis. Our models indicated that prey choice and consumption reflected seasonal changes in leopard seal population size and structure, size-selective predation and prey vulnerability. A population of 104 (90–125) leopard seals, of which 64% were juveniles, consumed less than 2% of the Antarctic fur seal pup production of the area (50% of total ingested energy, IE), but ca. 12–16% of the local gentoo penguin population (20% IE). Antarctic krill (28% IE) were the only observed food of leopard seal pups and supplemented the diet of older individuals. Direct impacts on krill and fish were negligible, but the “escapement” due to leopard seal predation on fur seal pups and penguins could be significant for the mackerel icefish fishery at South Georgia. These results suggest that: (1) rare apex predators like leopard seals may control, and may depend on, populations of mesopredators dependent on prey species targeted by fisheries; and (2) predatory impacts and community control may vary throughout the predator's geographic range, and differ across ecosystems and management areas, depending on the seasonal abundance of the prey and the predator's dispersal movements. This understanding is important to integrate the predator needs as natural mortality of its prey in models to set prey catch limits for fisheries. Reliable estimates of the variability of these needs are essential for a precautionary interpretation in the context of an ecosystem-based management.     

Geospatial variability of krill and top predators within an Antarctic submarine canyon system

The spatial distribution patterns of krill, seabirds (penguin, petrel and albatross), fur seals and baleen whales were mapped in nearshore waters (<50 km from land) to investigate their habitat selection within two adjacent submarine canyons near Livingston Island, Antarctica. Three shipboard surveys were conducted (February 2005–2007), and an echosounder was used to measure the distribution and abundance of krill while simultaneously conducting visual surveys to map seabird and marine mammals. Using a multispecies approach, we test the hypothesis that spatial organization of krill and top predators co-vary according to fine-scale changes in bathymetry in the nearshore marine environment. GAMs are used to examine the effect of sea depth, slope and distance to isobaths on the spatial distribution and abundance of krill and predators. Spatial distribution patterns of krill and predators relate to fine-scale (1–10 km) changes in bathymetry and exhibit cross-shelf gradients in abundance. Krill were concentrated along the shelf-break and abundant within both submarine canyons. Predators exhibited different preferences for locations within the submarine canyon system that relates to their foraging behavior. Penguins concentrated closer to shore and within the head of the east submarine canyon immediately adjacent to a breeding colony. Whales were also concentrated over the head of the east canyon (overlapping with penguins), whereas albatrosses and fur seals were concentrated in the west canyon. Fur seals also showed preference for steep slopes and were concentrated along the shelf-break. Petrels exhibited peaks in abundance throughout both submarine canyons. Owing to their orientation, size and proximity to the coastline, submarine canyons provide important habitat heterogeneity for krill and a variety of predators. This study highlights the multispecies approach for studying spatial ecology of top predators and krill and has implications for marine spatial management of the Scotia Sea.     

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.