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.     

Short- and long-term consistency in the foraging niche of wandering albatrosses

The wandering albatross (Diomedea exulans) is regarded as a generalist predator, but can it be consistent in its foraging niche at an individual level? This study tested short- and long-term consistency in the foraging niche in terms of habitat use, trophic level and, by inference, prey selection. Fieldwork was carried out at Bird Island, South Georgia, in May–October 2009, during the chick-rearing period. Blood (plasma and cells) and feathers for stable isotope analyses (δ¹³C and δ¹⁵N) were sampled from 35 adults on their return from a foraging trip during which they carried stomach temperature, activity and global positioning system loggers. Results suggest short-term consistency in foraging niche in relation to both oceanic water mass and trophic level, and long-term consistency in use of habitat. Consistent differences between individuals partly reflected sex-specific habitat preferences. The proportion of consistent individuals (i.e., with a narrow foraging niche) was estimated at c. 40?% for short-term habitat and trophic level (prey) preferences and 29?% for longer-term habitat preference, suggesting this is an important characteristic of this population and potentially of pelagic seabirds in general. Foraging consistency was not related to body condition or level of breeding experience; instead, it may reduce intraspecific competition.     

Albatross populations in peril: a population trajectory for black-browed albatrosses at south Georgia.

Simulation modeling was used to reconstruct Black-browed Albatross (Diomedea melanophris) population trends. Close approximations to observed data were accomplished by annually varying survival rates, reproductive success, and probabilities of returning to breed given success in previous years. The temporal shift in annual values coincided with the start of longline fishing at South Georgia and potential changes in krill abundance. We used 23 years of demographic data from long-term studies of a breeding colony of this species at Bird Island, South Georgia, to validate our model. When we used annual parameter estimates for survival, reproductive success, and probabilities of returning to breed given success in previous years, our model trajectory closely followed the observed changes in breeding population size over time. Population growth rate was below replacement (lambda < 1) in most years and was most sensitive to changes in adult survival. This supports the recent IUCN uplisting of this species from "Vulnerable" to "Endangered." Comparison of pre-1988 and post-1988 demography (before and after the inception of a longline fishery in the breeding area) reveals a decrease in lambda from 0.963 to 0.910. A life table response experiment (LTRE) showed that this decline in lambda was caused mostly by declines in survival of adults. If 1988-1998 demographic rates are maintained, the model predicts a 98% chance of a population of fewer than 25 pairs within 78 years. For this population to recover to a status under which it could be "delisted," a 10% increase in survival of all age classes would be needed.     

Growth of Antarctic krill Euphausia superba at South Georgia

 Antarctic krill Euphausia superba has a central role in the ecosystem of the Southern Ocean and knowledge of its growth rate is central to determining the factors influencing population dynamics. The length of Antarctic krill in the diet of Antarctic fur seals Arctocephalus gazella at South Georgia revealed a consistent increase in size between ca. 42 and ca. 54 mm over the period October–March, indicating growth rates much higher than predicted by existing models. Geographical variation in growth rate may result in 2-year-old krill at South Georgia attaining the same size as 3-year-old krill in the Antarctic Peninsula region. The effect of geographical variation in growth rate on the population structure of krill has important implications for comparing the fate of individual cohorts over large scales and in the interpretation of krill life-cycles. Received: 20 May 2000 / Accepted: 11 August 2000     

Inter-annual variation in the cephalopod component of the diet of the wandering albatross, <Emphasis Type="Italic">Diomedea exulans</Emphasis>, breeding at Bird Island,South Georgia

Cephalopods play an important role in the diet of many predators of the Southern Ocean. We investigated the cephalopod component of the diet of the wandering albatross, during breeding at South Georgia, from 269 boluses containing more than 34,000 beaks (corresponding to 19,452 individual cephalopods), collected between 1989 and 1999, and assessed the inter-annual variability of the cephalopod species, the level of scavenging on cephalopods by wandering albatrosses and the relationships between cephalopod availability and wandering albatross breeding parameters (breeding period, breeding success, fledging period, fledging success, egg mass and chick mass). We also proposed possible foraging areas of wandering albatrosses based on the cephalopods eaten. The cephalopod component of the diet of wandering albatrosses was relatively stable over the 11 years of the study. By number of lower beaks, three species predominated in the cephalopod component of the diet: Kondakovia longimana (29.5% of the total number of lower beaks; family Onychoteuthidae), Taonius sp. (20.4%; family Cranchiidae) and Histioteuthis sp. B (19.5%; family Histioteuthidae). K. longimana was consistently the predominant cephalopod species by mass in every year of the study (range 60.2-88.7% of mass contribution to diet). A minimum estimate of 30% of the number of cephalopods scavenged corresponded to 85% of the total mass of the cephalopods that contributed to the diet. Wandering albatrosses fed consistently more on "Antarctic" cephalopods than on "sub-Antarctic" or "subtropical" cephalopods in all years of the study, suggesting that Antarctic waters are an important foraging area for wandering albatrosses. Although some significant correlations between cephalopod abundance and wandering albatross breeding parameters existed (e.g. correlation between Taonius sp. by number and fledging success), none included K. longimana. When comparing groups of variables (using canonical analysis), no correlations were found between the most important cephalopod species (by number and by mass), total mass of squid consumed, cephalopod diversity index (H) for each year, and wandering albatross breeding parameters. This may reflect the possibility that other components in the diet (e.g. fish and carrion) are more important or, more likely, that the consistency across years of the wandering albatross breeding performance indicates that it is well buffered against fluctuations in prey availability.     

Feeding rates and diel vertical migration of copepods near South Georgia: comparison of shelf and oceanic sites

Seventeen Longhurst Hardy Plankton Recorder profiles were taken over a diel cycle in January 1990 to study the feeding of four major copepods over the South Georgia shelf. Ontogenetic changes in vertical migration were followed and feeding cycles determined by gut fluorometry for Calanoides acutus Stage CV, Calanus sinillimus CV and CVI, C. propinquus CV and Rhincalanus gigas CV and CVI. In common with a neighbouring oceanic site visited two weeks later and reported elsewhere, all four species had a diel cycle of feeding and migration. The vertical distributions of C. simillimus (all stages), R. gigas (nauplii) and Euphausia frigida (postlarvae) were similar at both sites, the night being spent within the chlorophyll maximum at 15 to 30 m. However, the biomass dominants, C. acutus and R. gigas, dwelt below the chlorophyll maximum, about 30 m deeper than their oceanic counterparts. Unlike the oceanic site, feeding at the shelf site was not restricted to darkness, but increased 6 to 10 h before nightfall and finished at dawn; the intervening period coincided with sinking and digestion. Daylight feeding may have been induced by the shorter night, lower light levels or greater food requirements at the shelf site, despite planktonic predators being over three times more abundant. Daily ration estimates for R. gigas at both sites were only 2% body carbon per day. These low values contrast with its smaller competirors, whose rations were in the range 5.6 to 27%.     

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.     

Diel vertical migration and feeding of copepods at an oceanic site near South Georgia

Twelve Longhurst Hardy Plankton Recorder (LHPR) profiles were taken over a 16 h period in January 1990, in order to study feeding of four copepod species at an Antarctic oceanic site near South Georgia. Vertical distributions of their life stages, as well as those of dominant competitors and predators, are described in relation to the feeding cycles of Calanoides acutus CV, Calanus simillimus CV, Calanus propinquus CV and Rhincalanus gigas CIII, CV and CVI. Comparisons with vertical ring-net catches, which were used for concomitant gutevacuation experiments, demonstrated the suitability of the LHPR for these fine-scale studies. Planktonic predators, with the exception of the diel migrant Themisto gaudichaudii, resided deeper than the herbivores. During the day and around midnight, when feeding rates were low, species and stages reached their maximum vertical separation. At these times, new generation copepodites of the four species lived progressively deeper and the overwintered generation (i.e., R. gigas Stages CIV, CV, CVI) were progressively shallower. During the afternoon or evening (depending on species), all stages older than CII, as well as Euphausia frigida and T. gaudichaudii, migrated upwards, to amass in the surface mixed layer. Feeding was restricted to darkness, although R. gigas commenced several hours before dusk. In detail their migration and feeding differed widely, with combinations of unimodal and apparent bimodal cycles. As a whole, the results suggest that (1) feeding could occur during sinking as well as during upward migrations, (2) upward migrations were not always associated with feeding increases, and (3) individuals appeared to descend after filling their guts.     

Predicting changes in the Antarctic krill, Euphausia superba, population at South Georgia

Variability in the Southern Ocean is frequently reflected in changes in the abundance of Antarctic krill Euphausia superba and subsequent effects on dependent predators. However, the nature and consequences of changes in krill population dynamics that accompany fluctuations in its abundance are essentially unknown. A conceptual model, developed from quantitative measures of krill length in the diet of predators at South Georgia from 1991 to 1997, allowed predictions to be made about the abundance and population structure of krill in 1998 and the consequences for predators. Consistent with model predictions, in 1998 there was a serial change in krill population structure, low krill biomass and low reproductive performance of predators. The change in the modal size of krill, from 56 mm in December to 42 mm in March, was apparently a result of the transport of krill into the region. This is the first occasion when the future status and structure of the krill population at South Georgia has been successfully predicted. By representing local krill population dynamics, which may also reflect large-scale physical and biological processes, predators have a potential key role as indicators of environmental variation in the Southern Ocean at a range of spatial scales. Received: 6 March 1999 / Accepted: 3 September 1999     

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.     

Upwelling-generated plankton strandlines: important predictable food sources for seabirds at Husvik,South Georgia

Sustained strong offshore winds generate upwelling in the fjords of northern South Georgia. Deepwater plankton is pumped by upwelling towards sandy beaches at the heads of the fjords; calanoid copepods, chaetognaths and pteropods are stranded by the sand filter in large quantities on falling tides. Local benthonic species (harpacticoid copepods and amphipods) are present in the plankton but do not strand. Strandlines are very rich (<4 kg wet mass m^-2) and provide large fractions of the food of kelp gulls, sheathbills and terns in the austral summer. Birds can only exploit fresh strandlines, since air-drying of the plankton soon makes it too salty to eat.     

Use of radar detectors to track attendance of albatrosses at fishing vessels

Despite international waters covering over 60% of the world's oceans, understanding of how fisheries in these regions shape ecosystem processes is surprisingly poor. Seabirds forage at fishing vessels, which has potentially deleterious effects for their population, but the extent of overlap and behavior in relation to ships is poorly known. Using novel biologging devices, which detect radar emissions and record the position of boats and seabirds, we measured the true extent of the overlap between seabirds and fishing vessels and generated estimates of the intensity of fishing and distribution of vessels in international waters. During breeding, wandering albatrosses (Diomedea exulans) from the Crozet Islands patrolled an area of over 10 million km² at distances up to 2500 km from the colony. Up to 79.5% of loggers attached to birds detected vessels. The extent of overlap between albatrosses and fisheries has widespread implications for bycatch risk in seabirds and reveals the areas of intense fishing throughout the ocean. We suggest that seabirds equipped with radar detectors are excellent monitors of the presence of vessels in the Southern Ocean and offer a new way to monitor the presence of illegal fisheries and to better understand the impact of fisheries on seabirds.     

At-sea activity patterns of breeding and nonbreeding white-chinned petrels <Emphasis Type="Italic">Procellaria aequinoctialis</Emphasis> from South Georgia

Despite the recent burgeoning in predator tracking studies, few report on seabird activity patterns, despite the potential to provide important insights into foraging ecology and distribution. In the first year-round study for any small petrel, we examined the activity patterns of the white-chinned petrel Procellaria aequinoctialis based on data from combination geolocator-immersion loggers deployed on adults at South Georgia. The petrels were highly nocturnal, flying for greater proportions of darkness than any large procellarid studied so far, except the light-mantled albatross Phoebetria palpebrata. Flight bout durations were short compared with other species, suggesting a dominant foraging mode of small-scale searching within large prey patches. When migrating, birds reduced the proportion of time on the water and increased flight bout duration. Activity patterns changed seasonally: birds flew least during the nonbreeding period, and most frequently during chick-rearing in order to meet higher energy demands associated with provisioning offspring. The degree of their response to moonlight was also stage dependent (greatest in nonbreeding, and weakest in incubating birds), a trait potentially shared by other nocturnal petrels which will have repercussions for feeding success and prey selection. For the white-chinned petrel, which is commonly caught in longline fisheries, these results can be used to identify periods when birds are most susceptible to bycatch, and therefore when use of mitigation and checking for compliance is critical.     

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.     

Feeding ecology of the green turtle (Chelonia mydas) at rocky reefs in western South Atlantic

Feeding ecology of green turtles was investigated between January 2005 and April 2008 at Arvoredo Reserve, Brazil (27°17′S, 48°18′W). Data were obtained through the performance of observational sessions, geo-referenced counts, benthic surveys, capture and recapture of individuals, and oesophageal lavages. This protected area was identified as an important green turtle feeding ground, used year-round by juveniles (curved carapace length = 32–83 cm). Turtles fed close to the rocky shores of the area and selected grazing sites commonly at hard-to-reach, near-vertical portions of the rocks. They were less active in cold months, and more abundant at shallow areas of the reef (0–5 m), where their preferred food items occurred. Their diet was dominated by macroalgae species but invertebrates were also present. Their main food item was the red algae Pterocladiella capillacea, which seems to be eaten through periodical cropping of its tips. Observational methods such as the ones applied here could be incorporated to other research programs aiming to understand the relationships between Chelonia mydas feeding populations and their environment.     

Biochemical composition and trophic strategies of the amphipod Eurythenes gryllus at hadal depths (Atacama Trench,South Pacific)

Amphipods Eurythenes gryllus were collected at 7800?m depth in the Atacama Trench (South Pacific) for studying their biochemical composition (in terms of proteins, lipids and carbohydrates and fatty acid content) and to gather information on bioenergetic strategies and trophic habits of organisms living in this extreme environment. The effect of long-term formalin storage on the biochemical determinations was also determined. Proteins were the dominant biochemical class of organic compounds (39–53%D.W.), whereas carbohydrates accounted for a very small fraction (1–2%D.W.). Lipid concentrations of E. gryllus accounted for 7–18%D.W. and were much lower than those reported for other deep-sea amphipods. These differences are likely to be more dependent upon food availability in the Atacama Trench rather than upon temperature. Lipid composition of E. gryllus revealed the dominance of monounsaturated fatty acids with polyunsaturated fatty acids accounting for a very small fraction, suggesting that hadal amphipods are higher dependent upon lipid reserves than amphipods inhabiting at shallow depths. The ratio of C18:1Δ⁹ to C18:1Δ¹¹ was >11 confirming the necrophagous trophic habits of this hadal amphipod.     

Distribution and diet of juvenile Patagonian toothfish on the South Georgia and Shag Rocks shelves (Southern Ocean)

The distribution and diet of juvenile (<750 mm) Patagonian toothfish are described from four annual trawl surveys (2003–2006) around the island of South Georgia in the Atlantic sector of the Southern Ocean. Recruitment of toothfish varies inter-annually, and a single large cohort dominated during the four years surveyed. Most juveniles were caught on the Shag Rocks shelf to the NW of South Georgia, with fish subsequently dispersing to deeper water around both the South Georgia and Shag Rocks shelves. Mean size of juvenile toothfish increased with depth of capture. Stomach contents analysis was conducted on 795 fish that contained food remains and revealed that juvenile toothfish are essentially piscivorous, with the diet dominated by notothenid fish. The yellow-finned notothen, Patagonotothen guntheri, was the dominant prey at Shag Rocks whilst at South Georgia, where P. guntheri is absent, the dominant prey were Antarctic krill and notothenid fish. The diet changed with size, with an increase in myctophid fish and krill as toothfish grow and disperse. The size of prey also increased with fish size, with a greater range of prey sizes consumed by larger fish.     

Comparison of light- and SST-based geolocation with satellite telemetry in free-ranging albatrosses

Light-based archival tags are increasingly being used on free-ranging marine vertebrates to study their movements using geolocation estimates. These methods use algorithms that incorporate threshold light techniques to determine longitude and latitude. More recently, researchers have begun using sea surface temperature (SST) to determine latitude in temperate regions. The accuracy and application of these algorithms have not been validated on free-ranging birds. Errors in both geolocation methods were quantified by double-tagging Laysan (Phoebastria immutabilis Rothschild) and black-footed (P. nigripes Audubon) albatrosses with both leg-mounted archival tags that measured SST and ambient light, and satellite transmitters. Laysan albatrosses were captured and released from breeding colonies on Tern Island, northwestern Hawaiian Islands (23°52N, 166°17W) and Guadalupe Island, Mexico (28°31N, 118°10W) and black-footed albatrosses from Tern Island. Studies were carried out between December 2002 and March 2003. For all birds combined, the mean ± SD great circle (GC) distance between light-based locations and satellite-derived locations was 400±298 km (n=131). Errors in geolocation positions were reduced to 202±171 km (n=154) when light-based longitude and SST-based latitude (i.e. SST/light) were used to establish locations. The SST/light method produced comparable results for two Laysan albatross populations that traveled within distinctly different oceanic regions (open ocean vs more coastal) whereas light-based methods produced greater errors in the coastal population. Archival tags deployed on black-footed albatrosses returned a significantly higher proportion of lower-quality locations, which was attributed to interference of the light sensor on the tag. Overall, the results demonstrate that combining measures of light-based longitude and SST-based latitude significantly reduces the error in location estimates for albatrosses and can provide valid latitude estimates during the equinoxes, when light-based latitude measurements are indeterminate.Communicated by J.P. Grassle, New Brunswick     

Abundance,breeding and growth of the crab Scylla serrata in two South African estuaries

Populations of the crab Scylla serrata (Forskal) were studied in two South African estuaries from 1971 until 1974. One estuary was open to the sea, the other was closed by a sandbar. Population density in the closed estuary was estimated by means of mark-release-recapture data from tagged crabs. The population of S. serrata was estimated as 1 crab/124m² and the production at 3.4 g/m²/annum. Catch per unit effort data from the closed estuary indicated a natural mortality of 41% in the crabs' second year and 60% in the third. Growth was studied in both estuaries by means of size-frequency analysis and from tagging returns. Growth was rapid in the first 12 to 15 months, when the crabs attained a carapace width of 80 to 160 mm. Thereafter growth slowed, and after 3 years crabs were between 140 and 180 mm in carapace width. Females mated in summer at a carapace width of 103 to 148 mm, males at 141 to 166 mm. After mating, females migrated out of both estuaries into the sea.