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     

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.     

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.     

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.     

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.     

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     

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.     

High male reproductive success in a low-density Antarctic fur seal (Arctocephalus gazella) breeding colony

Understanding how population density influences mating systems may lead to important insights into the plasticity of breeding behavior, but few natural systems allow for such studies. Antarctic fur seals (Arctocephalus gazella) provide an interesting model system because they breed in colonies of varying densities. Previous studies have largely focused on a high-density site at Bird Island, South Georgia. Here, 13 highly polymorphic microsatellite loci were used to conduct a genetic analysis of a low-density breeding colony of this species at Livingston Island, approximately 1,600 km south of South Georgia. The majority of adults seen ashore (n?=?54) were sampled together with every pup born (n?=?97) over four consecutive years. Paternities were confidently assigned for 34 out of the 97 pups. Two out of 23 sampled males accounted for the paternity of 28 % of all pups sampled during the study and 82 % of the pups with an assigned father. Moreover, a full likelihood pedigree inference method assigned a further eight paternities to a single unsampled male seal that is inferred to have held a territory during the season before the study began. The most successful males in our study easily surpassed the previous record for the total number of pups sired per male seal for the species. Furthermore, we identified two triads of full siblings implying that their parents remated in three consecutive years. These findings suggest that territorial male fur seals may achieve greater success in monopolizing access to breeding females when population density is relatively low.     

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.     

Foraging behaviour and habitat use by brown skuas Stercorarius lonnbergi breeding at South Georgia

Top predators are critical to ecosystem function, exerting a stabilising effect on the food web. Brown skuas are opportunistic predators and scavengers. Although skuas are often the dominant land-based predator at seabird colonies, this is the first detailed study of their movements and activity during breeding. The study was carried out at Bird Island, South Georgia (54°00′S, 38°03′W), in the austral summer of 2011/2012 and included GPS data from 33 breeding adults tracked during the late incubation and early chick-rearing periods. Brown skuas spent on average more than 80 % of time in the territory, and it was extremely rare for both partners to leave the territory simultaneously. Much more time was spent foraging at the coast than in penguin colonies and, based on saltwater immersion data, adults never foraged at sea. None of the tracked birds appeared to specialise in catching small petrels at night. Fewer foraging trips were made per day, and hence, more time was spent in the territory, during incubation than chick-rearing. Despite the pronounced sexual size dimorphism, there were no effects of sex on territorial attendance, foraging time or habitat use. Skuas at Bird Island show higher territorial attendance and are less likely to leave the territory unattended than those breeding elsewhere, suggesting closer proximity to more diverse or abundant food resources than at other colonies. The results tie in with previous diet studies, indicating that brown skuas at this site feed mostly on seal placentae and carrion and that birds may rely on a broader range of food resources as the season progresses.     

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.     

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.     

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.     

Activity patterns of giant petrels, Macronectes spp., using different foraging strategies

We studied foraging activity of giant petrels during the incubation period, by simultaneously deploying activity recorders and satellite transmitters on northern (Macronectes halli) and southern giant petrels (Macronectes giganteus) at Bird Island (South Georgia, Antarctica) between 29 October and 26 December 1998. Satellite tracking showed two types of trips: (1) coastal trips, all undertaken by male northern giant petrels, to the nearby South Georgia mainland, presumably foraging on seal and penguin carcasses on beaches, and (2) pelagic trips, foraging at sea for marine prey or potentially scavenging on distant archipelagos (e.g. South Sandwich, Falkland or South Orkney Islands). Activity recorder data were consistent with the types of trip defined by the satellite tracking data, with median wet activity (time spent at the sea surface) during pelagic trips being 41%, but only 14% on coastal trips. On pelagic trips, there was a significant negative correlation between the duration of wet periods and the speed of travel between satellite uplinks. Mean travelling speed between uplinks was greater during day than night for both types of trips, suggesting that giant petrels prefer to travel during daylight and are less active at night. The scarcity of wet periods during the night in giant petrels foraging to the South Georgia coast (median=3%, range=1-9%) indicates that such birds spent almost all night on land. Likewise, the scarcity of wet periods at night for three birds foraging 700-1,000 km south of Bird Island, where there is no land but abundant icebergs, suggests these birds were resting on the icebergs at night. In addition to the adaptations to scavenging on carrion, pelagic trips by giant petrels contain elements similar to those of albatrosses, indicating a complexity to giant petrel lifestyle hitherto unrecognised.     

Impact of miniature geolocation loggers on a small petrel, the thin-billed prion Pachyptila belcheri

Effects of deployment of miniaturised transmitters and loggers have been studied mainly in diving seabirds such as penguins, and less so in flying seabirds. However, some studies of albatrosses and petrels recorded extended trip durations and elevated rates of nest desertion following device attachment, especially if transmitter loads exceeded 3 % of adult mass. Studies have usually compared performance parameters such as trip duration, meal mass, breeding success or rate of return in the next season between birds with devices and controls. We here examined the effects of geolocator loggers (Global Location Sensing, (GLS)) on thin-billed prions Pachyptila belcheri (130 g), by comparing performance parameters and additionally eco-physiological parameters. GLS weighed ca. 1 % of the body mass, and were fixed on leg rings, which may influence the flight efficiency by creating an asymmetric load. We found no differences in the performance parameters, either in the season of attachment or the season following recovery. Similar stable isotope ratios in adult blood and feather samples further indicated that the foraging ecology was not influenced. However, after 1 year of logger deployment, adults differed in their hormonal response to stress: while baseline corticosterone levels were not influenced, corticosterone levels in response to handling were elevated. Moreover, increased heterophil/lymphocyte ratios and a decreased tail growth in winter suggest that carrying the GLS was energetically costly, and adults adapted physiologically to the higher work load, while keeping up a normal breeding performance.     

Feeding strategies and diets of breeding grey-headed and wandering albatrosses at South Georgia

The foraging areas and diets of the grey-headed albatross Thalassarche chrysostoma and wandering albatross Diomedea exulans were studied in March/April 2000 at Bird Island, South Georgia, during their respective chick-rearing and brood-guard periods. Oceanographically, March/April 2000 was abnormal, with warm conditions close to South Georgia. These conditions affected albatross foraging behaviour, particularly that of grey-headed albatrosses. Both species tended to forage in different areas of the ocean, with significant differences in trip durations. Grey-headed albatrosses (n=9) foraged mainly in Antarctic waters (predominantly shelf waters of the South Shetland Islands and Antarctic Peninsula, and also in oceanic waters around South Georgia), feeding mainly on krill (Euphausia superba; 77% by mass). Foraging trips lasted 13.3 days (range: 5–26 days), far longer than the 1–3 days found in previous studies. Only one grey-headed albatross was associated with the APF (Antarctic Polar Front), a reported foraging area in recent studies. Wandering albatrosses (n=9) foraged in Antarctic (South Georgia Shelf) and Antarctic Polar Frontal Zone (APFZ) waters, with trips of 1–4 days trip duration (usual for this species), feeding on fish (46% by mass) and cephalopods (32%). One bird was associated with the APF, and two birds foraged on the shelf/shelf break over the Patagonian shelf. These findings suggest that sea surface temperature anomalies, produced by movement of the APF closer to South Georgia or by eddies, may have had an effect on the foraging strategy of grey-headed albatrosses that year (the main prey of grey-headed albatrosses in previous studies, the ommastrephid Martialia hyadesi, is known to be associated with the APF). Also, when both albatross breeding periods overlap, their foraging areas were complementary, which reflected the prey taken.     

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.     

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.     

A method to estimate pre-exploitation population size

Antarctic fur seals (Arctocephalus gazella) were commercially exploited on the subantarctic island of South Georgia for over 100 years and nearly driven to extinction. Since the cessation of harvesting, however, their populations have rebounded, and they are now often considered a nuisance species whose impact on the terrestrial landscape should be mitigated. Any evaluation of their current population requires the context provided by their historic, pre-exploitation abundance, lest ecologists fall prey to shifting baseline syndrome in which their perspective on current abundance is compared only with an altered state resulting from past anthropogenic disturbance. Estimating pre-exploitation abundance is critical to defining species recovery and setting recovery targets, both of which are needed for the International Union for the Conservation of Nature's recent efforts to develop a green list of recovering species. To address this issue, we reconstructed the South Georgia fur seal harvest from 1786 to 1908 from ship logbooks and other historical records and interpolated missing harvest data as necessary with a generalized linear model fit to the historical record. Using an approximate Bayesian computation framework, harvest data, and a stochastic age-structured population model, we estimated the pre-exploitation abundance of Antarctic fur seals on South Georgia was 2.5 million females (95% CI 1.5–3.5 million). This estimate is similar to recent abundance estimates, and suggests current populations, and the ecological consequences of so many fur seals on the island, may be similar to conditions prior to human harvest. Although the historic archive on the fur sealing era is unavoidably patchy, the use of archival records is essential for reconstructing the past and, correspondingly, to understanding the present. Article impact statement: Defining species recovery requires an understanding of baseline population state, which can be estimated through statistical methods.     

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.