Combining DNA and morphological analyses of faecal samples improves insight into trophic interactions: a case study using a generalist predator

The diet of pinnipeds is most commonly inferred from morphologically diagnostic remains of prey in their scats. Although this method can generate quantitative estimates of diet simply, important prey types may not always be detected. DNA-based methods improve detection of prey in scats, but they are not quantitative. While some studies have combined morphological and DNA-based methods, these have only assessed prey that are represented by their hard remains in scats. To overcome this bias, we apply molecular and morphological analyses to the soft and hard portions of faecal samples respectively, to estimate the diet of lactating Antarctic fur seals (Arctocephalus gazella) on Heard Island. The diet of this population is of particular interest because it is expanding rapidly and may rely to some extent on mackerel icefish (Champsocephalus gunnari), which are subject to commercial fisheries. Based on results from morphological analysis and likely important prey types, we tested for DNA remains of C. gunnari, myctophids and squid in faecal samples. The proportion of samples (n = 54) yielding no dietary information was reduced from around 25.9% using either method alone, to 9.3% when combined. Detection of all prey types tested for was notably improved by integrating molecular and morphological data. Data from either method alone would have underestimated the number of animals consuming C. gunnari by around 25.7%. Detection of multiple prey types in samples increased from 9.3% when using morphological analysis only, to 33.3% when using DNA only, to 46.3% when using both methods. Taken in isolation, morphological data inferred that individual seals consume either C. gunnari or myctophids, probably foraging in separate locations characteristic of those prey. Including molecular data demonstrated that while this may be true of some individuals, many other seals consume a mixed diet of at least C. gunnari, myctophids and squid. This new approach of combining DNA-based and morphological analyses of diet samples markedly increased the number of samples yielding dietary information, as well as increasing the amount of information attained from those samples. Our findings illustrate the broad potential of this technique to improve insight into trophic interactions in marine ecosystems.

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A case of parasite-mediated competition? Phenotypic differentiation among hookworms Uncinaria sp. (Nematoda: Ancylostomatidae) in sympatric and allopatric populations of South American sea lions Otaria byronia,and fur seals Arctocephalus australis (Carnivora: Otariidae)

Hookworms in otariids are considered to meet the conditions and to fulfill the predictions set forth in the theoretical literature to define a likely case of host coexistence mediation by a shared pathogen. The intensity of infections, the prevalence of skin lesions and the morphometry of hookworms Uncinaria spp. were examined in South American sea lions and fur seals sampled along the Chilean and the Uruguayan coasts in spring and summer 1981 to 1991. In sympatric host populations from Uruguay, there were clear differences in intensity of infections, prevalence of skin lesions, and body size of the hookworms from the two host species. Sea lions from Chile, allopatric to fur seal populations, are less intensively infected and hookworms found in these sea lions have the smallest body size reported in otariids, while those from South American fur seals are the largest. Hookworms found in sea lion pups from populations sympatric to fur seals revealed intermediate values in intensity and in body size, and the sea lion pups had the highest prevalence of skin lesions. Other reports of hookworms in otariids show intermediate morphometry, following a general linear trend of differentiation in size. Consequently, they are considered to belong to the same, widely distributed species Uncinaria lucasi Stiles, 1901.     

Dynamic intra-seasonal habitat use by Antarctic fur seals suggests migratory hotspots near the Antarctic Peninsula

Antarctic fur seals (Arctocephalus gazella) are major secondary consumers in the Southern Ocean, placing them in potential competition with commercial fisheries and requiring research to understand their seasonal habitat use. Using the data obtained during 14 shipboard surveys sampled on a fixed grid (150 K km²) during mid- to late summer, I quantified the spatial distribution and intra-seasonal variability of fur seal sightings relative to distance to land and hydrographic boundaries. I test the hypothesis that fur seals display an increase in their at-sea abundance during mid- to late summer near the Antarctic Peninsula as they prepare to take up wintering grounds. I also test whether abundances of their potential prey, krill and myctophids, exhibit intra-seasonal variability. During midsummer, high-abundance areas are located near major breeding colonies; however, during late summer, there is an order-of-magnitude increase in fur seal abundance, coinciding with an increase in the number of high-abundance areas located in Bransfield Strait. Coincidently, abundance of Euphausia superba decreased and the myctophid Electrona antarctica increased between mid- and late-summer surveys. High-abundance areas of fur seals are not associated with the southern Antarctic Circumpolar Current front but are concentrated within 100 km from land, potentially indicating the location of haul out and important coastal habitat use areas. The dynamic increase in the number and location of high-abundance areas during late summer represents a considerable amount of mammalian predators entering the Antarctic Peninsula marine ecosystem. This information is important for understanding the seasonal impact of fur seals on regional marine food webs and their potential interaction with the autumn–winter krill fishery.     

Alimentary niche partitioning in the Galapagos sea lion, Zalophus wollebaeki

Sea lions are generally considered opportunistic feeders. However, studies from different areas suggest their diet consists mostly of four to five types of prey. Previous studies in Galapagos sea lions have identified at least three feeding strategies for this species, suggesting diversification of their diet. Diet diversification is favored in organisms with relatively high trophic position and subject to high intra-specific and low inter-specific competition. Zalophus wollebaeki meet these criteria as the only pinniped on San Cristobal Island, where three sea lion rookeries are located within 11 km: a distance considerably shorter than their 41 km foraging range. To measure the degree of diet diversification, we used scats and stable isotope analyses. A total of 270 scat samples from lactating females and 142 fur samples from sea lion pups were collected during the breeding season 2006. The scat analysis identified distinct diets among rookeries, with minimal trophic overlap (Cλ = 0.19), a trophic level TL = 4.5 (secondary–tertiary carnivore), and trophic breadth of a specialist predator (B _i  = 0.37). The mean δ¹⁵N and δ¹³C values were 13.07 ± 0.52 and −16.34 ± 0.37, respectively. No significant difference was found in the δ¹⁵N values from the sea lion rookeries, but differences were found inter- and intra-population in δ¹³C values for pups from different groups (ANOVA P < 0.05). Our results indicate that diet diversification is present in the Galapagos sea lion and may play important role to the survival of the species in a habitat where pinniped populations are limited.     

Spatial and temporal variation in the diet of a high trophic level predator,the Australian fur seal (<Emphasis Type="Italic">Arctocephalus pusillus doriferus</Emphasis>)

This study quantifies the manner in which Australian fur seals, Arctocephalus pusillus doriferus, use their prey in a spatial and temporal context. We analysed 977 scat and 66 regurgitate samples collected from Tasmanian breeding colonies and haul-outs between 1994 and 2000. Diagnostic prey remains identified in the scats represented 35 fish taxa and 8 cephalopod taxa. The main taxa identified in scats, where frequency of occurrence was 10%, were leatherjacket species (family Monocanthidae), redbait (Emmelichthys nitidus), barracouta (Thyrsites atun), jack mackerel (Trachurus declivis) and red cod (Pseudophysis bachus). Regurgitates were dominated by cephalopods, primarily Goulds squid (Nototodarus gouldi), Octopus maorum, O. berrima/pallidus and Sepia apama. Discriminant function analyses indicated that there were generally no significant differences in the composition of the diet between colonies within a year, suggesting that prey distribution is fairly uniform throughout Bass Strait at those time scales. The diet at breeding colonies, however, exhibited significant inter- and intra-annual variation, determined by the presence of several key taxa, such as barracouta and a species of scorpionfish (family Scorpaenidae). The diet composition also varied regionally, between Bass Strait and southern Tasmania in spring 1999 and autumn 2000, with redbait, barracouta and a species of scorpionfish identified as the main taxa contributing to this difference. Redbait occurred in the diet only in southern Tasmania, whereas barracouta and scorpionfish occurred only in Bass Strait.Electronic Supplementary Material Supplementary material is available in the online version of this article at http://dx.doi.org/10.1007/s00227-003-1219-0.Communicated by G.F. Humphrey, Sydney     

Temperature and salinity: two climate change stressors affecting early development of the New Zealand sea urchin Evechinus chloroticus

Temperature and salinity are important environmental factors affecting the normal functioning of marine animals, particularly animals such as sea urchins living in shallow waters and tide pools. Here, we evaluated the effect of different combinations of temperature and salinity on early embryos of the endemic New Zealand sea urchin Evechinus chloroticus. Animals were collected at Matheson’s Bay (36º18′17′′S; 174º47′51′′E) in north-eastern New Zealand in February 2013. Embryos were exposed to five salinities (29, 31, 34, 35 and 37 ppt) and two temperatures (18 and 21 °C) during the first 24 h of development. Low salinity (29 ppt) affected all parameters (fertilization, development rate, gastrulation and normal development), with ca. 50 % of embryos surviving at 29 ppt, whereas seawater temperature only affected development rate and gastrulation. An increase in temperature from 18 to 21 °C minimized the negative effect of low salinity (≤31 ppt) on development rate and gastrulation of E. chloroticus. Overall, the results of this study suggest that early embryos of E. chloroticus have developmental plasticity to withstand reductions in salinity up to 29 ppt; however, it is still unknown whether the surviving embryos will be able to complete larval development at low salinities, particularly whether the embryos and larvae are carried into extreme environments such as estuaries where salinity is even lower. Multistressor studies are very important for climate change research as multiple environmental factors will act together in the wild, having major consequences for development and recruitment of marine invertebrates.     

Male-biased sex ratios in New Zealand fur seal pups relative to environmental variation

A number of models have been proposed to provide adaptive explanations of sex-ratio variation in mammals. Two models have been applied commonly to primates and ungulates with varying success—the Trivers-Willard (TW) hypothesis, and the local resource competition (LRC) hypothesis. For polygynous, sexually dimorphic mammals, where males are larger and disperse more readily, these models predict opposite outcomes of sex-ratio adjustment within the same environmental context (high-resource years: TW—more sons; LRC—more daughters). However, many of the predictions of these two models can vary depending on factors influencing resource availability, such as environmental stochasticity, resource predictability, and population density. The New Zealand fur seal (Arctocephalus forsteri) is a polygynous mammal showing marked sexual dimorphism (larger males), with higher variation in male reproductive success expected. We provide clear evidence of male-biased sex ratios from a large sample of A. forsteri pups captured around South Island, New Zealand during 1996/1998, even after accounting for a sex bias in capture probability. The extent of the bias depended upon year and, in 1998, strong climatic perturbations (El Niño/Southern Oscillation, ENSO) probably reduced food availability. Significant male-biased sex ratios were found in all years; however, there was a significant decline in the male bias in 1998. There was no relationship between sex ratio and population density. We suggest that the sex-ratio bias resulted from the production of relatively more male pups. Under the density-independent scenario, the strong male bias in A. forsteri sex ratios is support for the TW model within an environment of high resource predictability. We suggest that some plasticity in the determination of pup sex among years is a mechanism by which A. forsteri females in New Zealand, and perhaps other otariid seals, can maximise fitness benefits when living in regions of high, yet apparently predictable, environmental variability. We also suggest that much of the inconsistency in the reported sex ratios for otariid seals results from the complex interaction of population density and environmental stochasticity influencing relative food availability over time.     

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.     

Diet of orange roughy,Hoplostethus atlanticus (Pisces: Trachichthyidae) on the Challenger Plateau,New Zealand

Stomach contents of 687 orange roughy [Hoplostethus atlanticus (Collett, 1889)] from the Challenger Plateau, sampled in March, July and November 1984, were examined. The relative importance of different components of the diet was assessed using three feeding indices which combine, in different ways, percentage frequency of occurrence, percentage number, and percentage weight of prey categories. For both sexes, for all size classes and at any time of the year, the most frequent and abundant components of the diet were prawns, followed by fish, squid, amphipods and mysids. The main natant decapod families were Oplophoridae, Pasiphaeidae, and Sergestidae. Fish, mainly Chauliodontidae and Myctophidae, were the most important prey by weight. Most prey species were benthopelagic and mesopelagic organisms which move towards the surface at night. However, H. atlanticus can be caught by bottom trawl (between 750 and 1 200 m) during any 24 h period, and there was no evidence of vertical migration in search of their prey. When the fish grow in length, there is a transition in the diet from prawns, mysids, and fish, to prawns, fish and squid. Squid were not found in the stomach contents of fish smaller than 20 cm. These dietary changes may be linked to modifications in morphology with growth. The relative proportions of the main dietary items, and in particular the natant decapod families, varied with time of the year. Also, the stomach data seem to indicate an increasing importance of fish and squid in the diet in deeper water. H. atlanticus appears to be an opportunistic predator, consuming a wide variety of invertebrates and fishes. Our results provide evidence to support the notion that benthopelagic predators which consume vertically migrating mesopelagic fish, have an important role in the transfer of energy to the benthos.     

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.     

Effects of the Presence of Official‐Looking Volunteers on Harassment of New Zealand Fur Seals

Abstract: An increased number of tourists viewing animals in the wild have increased stress on these animals (hereafter wildlife). Many wildlife‐viewing locations rely on voluntary compliance with posted regulations to protect animals from tourists because of the expense of employing on‐site enforcement personnel. Voluntary compliance, however, is ineffective. The presence of official‐looking volunteers may decrease the incidence of wildlife harassment by tourists. To test this possibility, we observed tourists interacting with 5‐ to 12‐month‐old New Zealand fur seals (Arctocephalus forsteri) at the popular Ohau Stream waterfall while in the absence or presence of a young woman in plain sight wearing a neon vest (i.e., observer) and when an observer was not present. We observed 254 tourist groups at the waterfall when young seals were present. The percentage of groups in which at least one person harassed (approached, touched, or threw objects) a young seal was two‐thirds lower when the official‐looking observer was present. Frequency of harassment was inversely related to observer presence. Programs in which volunteers work at tourist sites are popular in countries with high tourism rates, such as New Zealand. Our results show that a relatively inexpensive and effective tourism‐management strategy may be to post such volunteers as observers at sites where tourists view wildlife.     

Nutritional geometry and macronutrient variation in the diets of gannets: the challenges in marine field studies

Foraging theory proposes that the nutritional driver of food choice and foraging in carnivores is energy gain. In contrast, recent laboratory experiments have shown that several species of carnivore select prey that provides a diet with a specific balance of macronutrients, rather than the highest energy content. It remains, however, to be determined how nutritionally variable the foods of predators in the wild are, and whether they feed selectively from available prey to balance their diet. Here, we used a geometric method named the right-angled mixture triangle (RMT) for examining nutritional variability in the prey and selected diets of a group of wild carnivores and marine top predators, the gannets (Morus spp.). A prey-level diet analysis was performed on Australasian gannets (M. serrator) from two New Zealand locations, and the macronutrient composition of their chosen prey species was measured. We use RMT to extend the comparison in the compositions of foods and diets from Australasian gannets from Australia as well as Northern gannets (M. bassanus) and Cape gannets (M. capensis). We found nutritional variability at multiple scales: intra- and interspecific variability in the pelagic fish and squid prey themselves; and intra- and interspecific variability in the diets consumed by geographically disparate populations of gannets. This nutritional variability potentially presents these predatory seabirds with both opportunity to select an optimal diet, and constraint if prevented from securing an optimal diet.     

Parent–offspring and sibling conflict in Galápagos fur seals and sea lions

Parent–offspring conflict theory is well supported by theoretical arguments. However, empirical observations are often difficult to interpret and have contradicted one of its most appealing predictions that parent and offspring should disagree over killing of nest or littermates. We present the first examples of deadly conflict between siblings of different cohorts. In Galápagos fur seals (Arctocephalus galapagoensis) and sea lions (Zalophus wollebaeki), mothers often wean their single offspring at 2 years. This leads to a situation where up to 23% of all pups are born while the older sibling is still being nursed. Younger siblings are disadvantaged by being born lighter than neonates without older still dependent siblings. Pups born while an older sib is still dependent grow less in early life (fur seal) and suffer increased early mortality (both species) through direct aggression or scramble competition with the older sibling. This effect is much stronger in years of high sea surface temperature (El Niño) indicating low marine productivity and if the older offspring is a male. In both species, mothers interfere aggressively in this conflict by defending the younger offspring. In years of El Niño, intense resistance to maternal aggression by the older offspring happens frequently in the fur seal. Such resistance against weaning can induce maternal neglect of the newborn. Given substantial year to year variation in offspring growth, maternal aggression forces weaning in the older sibling only if it has reached sufficient size to support itself by foraging. In Galápagos fur seals, pups with older siblings can either represent insurance against loss of older offspring or extra reproductive value.     

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.     

Impacts of near future sea surface pH and temperature conditions on fertilisation and embryonic development in Centrostephanus rodgersii from northern New Zealand and northern New South Wales, Australia

Oceans are warming and becoming more acidic. While higher temperature and lower pH can have negative effects on fertilisation and development of marine invertebrates, warming may partially ameliorate the negative effect of lower pH. This study determined the effect of warming (3 °C) and decreased pH (0.3, 0.5, 1.1 units below ambient) on fertilisation and development in two populations of the sea urchin Centrostephanus rodgersii, one at its northern range limit (Coffs Harbour, New South Wales NSW, 30°27′S, 153°14′E) and the other one in New Zealand where the species may be a recent arrival (Mokohinau Islands, 35°56′S, 175°9′E). Both populations were sampled in August 2011. The two populations exhibited a differential response to temperature, while pH affected them similarly. Fertilisation was robust to pH levels forecast for 2100, and it was only slightly reduced at pH values forecast for 2300 (i.e. ≈5 and ≈8 % for the northern NSW and the New Zealand populations, respectively). Decreased pH (pH = 7.6) reduced the percentage of succeeding developmental stages. Progression through cleavage and hatching stages was faster at +3 °C in the New Zealand population but not in northern NSW urchins, while for the NSW population, there was a positive interaction between temperature and pH at hatching. Gastrulation was negatively affected by an extreme pH 7.0 treatment (60–80 % reduction) and least affected by increased temperature. The percentage of abnormal embryos at gastrulation increased significantly at +3 °C treatment in the northern NSW population. Predicted future increases in temperature may facilitate further expansion of the geographical range of C. rodgersii in New Zealand, with a minimal effect of concurrent reduced pH.     

Age-related shifts in the diet composition of southern elephant seals expand overall foraging niche

Southern elephant seals are important apex predators in a highly variable and unpredictable marine environment. In the presence of resource limitation, foraging behaviours evolve to reduce intra-specific competition increasing a species’ overall probability of successful foraging. We examined the diet of 141 (aged 1–3 years) juvenile southern elephant seals to test the hypotheses that differences between ages, sexes and seasons in diet structure occur. We described prey species composition for common squid and fish species and the mean size of cephalopod prey items for these age groups. Three cephalopod species dominated the stomach samples, Alluroteuthis antarcticus, Histioteuthis eltaninae and Slosarczykovia circumantarcticus. We found age-related differences in both species composition and size of larger prey species that probably relate to ontogenetic changes in diving ability and haul-out behaviour and prey availability. These changes in foraging behaviour and diet are hypothesised to reduce intra-specific food competition concomitant with the increase in foraging niche of growing juveniles.     

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.     

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.     

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.     

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.