Extrapair paternities in black-browed Thalassarche melanophris, grey-headed T. chrysostoma and wandering albatrosses Diomedea exulans at South Georgia

Extrapair paternities (EPP) are relatively common in passerines, but rare in seabirds. Like most seabirds, albatrosses are long lived, form long-term pair bonds and require biparental care for chick-rearing. Microsatellite analyses of 327 chicks from black-browed Thalassarche melanophris , grey-headed T. chrysostoma and wandering albatrosses Diomedea exulans over two breeding seasons revealed the presence of EPP in all three species. Though EPP rates varied between species and years, up to 21% of offspring were the result of extrapair matings. Rates were highest in wandering albatrosses (6–21%) followed by grey-headed (3–10%) and black-browed (0–9%) albatrosses. EPP rates were lower in 1998 compared to 1999 in both black-browed and grey-headed albatrosses, whereas the reverse was true for wandering albatrosses. Interspecific differences in EPP rates may reflect differences in breeding phenology and sexual size dimorphism. Differences in timing and frequency of breeding may promote different opportunities for interactions with birds other than their normal partner. The different breeding habitat, dispersion and mate-attraction rituals in wandering albatross, together with the disparity in size between the sexes may also offer more scope for higher rates of EPP. Despite extensive sampling within each colony, we were unable to identify sires for many of the extrapair young; however males from other colonies were involved, raising interesting questions regarding the timing and nature of such events.     

The fish diet of black-browed albatrossDiomedea melanophris and grey-headed albatrossD. Chrysostoma at South Georgia

 The fish component of the diet of black-browed and grey-headed albatrosses at South Georgia was investigated by intercepting 155 meals from adults arriving to feed chicks during February 1986 and 1994. Fish represented 30% and 72% by mass of the diet of black-browed albatrosses and 14% and 60% by mass of the diet of grey-headed albatrosses in 1986 and 1994 respectively. We determined the identity and quantified the contribution (by numbers, size and mass) of fish species mainly by using otoliths (54 representing 9 taxa and 57 representing 17 taxa in black-browed and grey-headed albatross samples respectively). For black-browed albatrosses in 1986 the main fish prey was Patagonotothen guntheri (77% of otoliths, 51% of estimated fish biomass) and a single large specimen of Icichthys australis (40% estimated biomass), whereas in 1994 Pseudochaenichthys georgianus was the main fish prey (57% of estimated biomass) with Magnisudis prionosa (30%) and Champsocephalus gunnari (12%) also making substantial contributions. Grey-headed albatross samples from 1986 were dominated by southern lampreys (40% by number, 79% of estimated biomass), lanternfish (32% of numbers, 9% by mass) and Patagonotothen guntheri (11% by mass); in 1994 Champsocephalus gunnari (42% by numbers, 24% by mass), Magnisudis prionosa (13% by number, 36% by mass), Muraenolepis microps (90% by number), Pseudochaenichthys georgianus (15% by mass) and lanternfish (18% by number but only 1% by mass) were the main prey. The importance of Patagonotothen guntheri to both species in 1986 and its absence in 1994 probably reflect albatrosses obtaining it from the commercial fishery, which was active in 1986 but closed in 1994. Otherwise the fish diet of black-browed albatrosses is dominated by krill-feeding fish, characteristic of the waters of the South Georgia shelf. In contrast, the grey-headed albatross diet comprises deeper water mesopelagic species, especially lanternfish, which reflect its affinity for the Antarctic Polar Frontal Zone and associated oceanic upwellings. Received: 28 June 1995 / Accepted: 8 October 1995     

Geographical variation in morphometry of black-browed and grey-headed albatrosses from four sites

Differences in morphometry between five populations of black-browed albatrosses (Diomedea melanophrys) and four populations of grey-headed albatrosses (D. chrysostoma) are examined. Two clear groups of black-browed albatrosses are evident, with birds from the subspecies Diomedea melanophrys impavida showing significant differences in several variables from those from the subspecies Diomedea melanophrys melanophrys. For groups from the latter subspecies, birds from South Georgia had larger measures than those from Kerguelen. A similar pattern to that of Diomedea melanophrys melanophrys was found between the groups of grey-headed albatrosses. Analysis of foraging distances relative to adult body-size index and the duration of chick-rearing periods suggests that differences in chick-provisioning rates between populations of conspecifics could account for at least some of the observed differences in adult morphometry. Accepted: 5 April 1999     

Comparison of census methods for black-browed albatrosses breeding at the Ildefonso Archipelago,Chile

A breeding population of black-browed albatrosses has been known to exist at the Ildefonso Archipelago, Chile, for >90 years but the population has never been censused using scientifically defendable methods. To estimate population size, and examine the accuracy and practicality of various census methods, the population was censused in the 2002/2003 breeding season using (a) ground-truthed aerial photography, (b) yacht-based photography, (c) ground counts, (d) quadrat sampling and (e) point-distance sampling. Compared to ground-truthed aerial photography (judged the most accurate) yacht-based photography underestimated population size by 55%, ground counts by 13%, quadrat sampling by 11% and point-distance sampling by 9%. Ground-truthed air photography revealed that in the 2002/2003 breeding season 47,000 pairs of black-browed albatrosses bred at the Ildefonso Archipelago. A repeat aerial census in 2006 suggested the size of the breeding population had not changed in the 4 years between the two censuses. After the Falkland Islands/Islas Malvinas, South Georgia and Diego Ramirez, the Ildefonso Archipelago holds the fourth largest population of black-browed albatrosses in the world.     

Cephalopod prey of the black-browed albatrossDiomedea melanophrys at South Georgia

Regurgitations were collected from 41 black-browed albatross adults feeding chicks at Bird Island (54°S 38°W), South Georgia in February 1986. The samples were sorted into recognisable food categories and weighed. Cephalopods were identified by means of the lower beak, or in some cases the gladius, and allometric equations were used to calculate mantle length and wet body weight represented by beaks. The samples contained 35.5%Euphausia superba, 30.9% cephalopods and 27.1% fish, by weight. A total of 21 samples contained recognisable cephalopod remains and 20 contained specimens that could be identified. In all, 50 cephalopod specimens, representing an estimated 6,866 g wet weight, were identified. The diet was dominated in terms of numbers, weight and percent occurrence by the ommastrephid squidMartialia hyadesi, and in most cases the entire squid was present with only partial digestion of the skin and arm armature. The cranchiid squidGaliteuthis glacialis was the only other cephalopod of numerical importance but no soft parts were present suggesting that, although significant in the diet of the adults, this species was not being fed to chicks. One specimen each ofGonatus antarcticus, Chiroteuthis sp.,Histioteuthis sp. B. and the octopodidPareledone polymorpha were also present. The cephalopod composition of the diet corresponded closely with a collection made 10 years earlier. The commonest species in the bird's diet,M. hyadesi, has not been found in net and jig samples at South Georgia although it has been taken from the Antarctic Polar Frontal Zone to the west of the Island. The presence of almost complete, undigested, specimens ofM. hyadesi in the bird's diet indicates that it occurs relatively close to South Georgia.M. hyadesi preys largely on myctophid fishes, which themselves prey on small zooplankters, so a significant component of the black-browed albatross diet depends on a food chain which largely by-passesE. superba.     

Global relationships amongst black-browed and grey-headed albatrosses: analysis of population structure using mitochondrial DNA and microsatellites

The population structure of black-browed (Thalassarche melanophris and T. impavida) and grey-headed (T. chrysostoma) albatrosses was examined using both mitochondrial DNA (mtDNA) and microsatellite analyses. mtDNA sequences from 73 black-browed and 50 grey-headed albatrosses were obtained from five island groups in the Southern Ocean. High levels of sequence divergence were found in both taxa (0.55-7.20% in black-browed albatrosses and 2.10-3.90% in grey-headed albatrosses). Black-browed albatrosses form three distinct groups: Falklands, Diego Ramirez/South Georgia/Kerguelen, and Campbell Island (T. impavida). T. melanophris from Campbell Island contain birds from each of the three groups, indicating high levels of mixture and hybridization. In contrast, grey-headed albatrosses form one globally panmictic population. Microsatellite analyses on a larger number of samples using seven highly variable markers found similar population structure to the mtDNA analyses in both black-browed and grey-headed albatrosses. Differences in population structure between these two very similar and closely related species could be the result of differences in foraging and dispersal patterns. Breeding black-browed albatrosses forage mainly over continental shelves and migrate to similar areas when not breeding. Grey-headed albatrosses forage mainly at frontal systems, travelling widely across oceanic habitats outside the breeding season. Genetic analyses support the current classification of T. impavida as being distinct from T. melanophris, but would also suggest splitting T. melanophris into two groups: Falkland Islands, and Diego Ramirez/South Georgia/Kerguelen.     

The Evangelistas Islets,Chile: a new breeding site for black-browed albatrosses

A previously undescribed population of black-browed albatrosses (Thalassarche melanophrys) is reported at the Evangelistas Islets, Straits of Magellan, Chile. The population was censused from aerial photographs taken on 13 October 2002 that yielded an estimate of the number of breeding pairs. A combined total of 4,670 pairs of black-browed albatrosses were found nesting at Elcano and Lobos Islets, 2 of the 4 islets in the Evangelistas group. This new record raises to four the number of breeding islands of this albatross species in Chile.     

The diet of black-browed albatrosses at the Diego Ramirez Islands,Chile

The diet of black-browed albatrosses was studied at Gonzalo Island, Diego Ramirez, Chile, during the early chick-rearing periods of 2000, 2001, and 2002. Diet composition was determined by sampling chick-stomach contents during January and February of each year. Reconstituted meal mass was similar throughout the study, with diet being dominated by fish in all 3 years. Overall, the main items taken were the fishes Macruronus magellanicus (66–89% by mass) and Micromesistius australis (2.6–3.7% by mass), which are both fisheries-related species, and the squid Martialia hyadesi (8–20% by mass). The distribution of the prey species indicates that black-browed albatrosses obtained the bulk of their food over the South American continental shelf, but also foraged at the Antarctic Polar Front. The prevalence in the diet of fish species discarded from fishing operations, and the presence of fish hooks and fish bait species, indicate a strong association with fisheries in southern Chile.     

A major autumn feeding ground for fin whales, southern fulmars and grey-headed albatrosses around the South Shetland Islands, Antarctica

The main aim of our long-term study on the at-sea distribution of the upper trophic levels in polar marine ecosystems is to deepen the understanding of the basic mechanisms affecting their distribution, i.e. hydrological factors such as water masses and fronts, pack ice and ice edge, eddies. A second goal consists in detecting spatial and temporal changes, with special attention to global climate changes, as well as possible consequences of (krill) fisheries. Seabirds and marine mammals were recorded during an autumn expedition of icebreaking RV Polarstern between Punta Arenas and the South Shetland Islands, Antarctica in March/ April 2012. During a total of 333 half-an-hour transect counts without width limitation, 113,500 seabirds were encountered, belonging to 40 species. The vast majority were southern fulmars and grey-headed albatross around the South Shetland Islands, with 76,800 and 7,000 individuals, respectively. Fin whale was by far the most abundant cetacean with 300 identified individuals in the same area, of which 100 in one count. These exceptional concentrations of fin whale seem to reflect an autumn pre-migration feeding aggregation. For most species, the majority was concentrated in very few counts, reflecting a very high patchiness and, as upper trophic levels, an important prey availability—mainly krill, nekton and small fish. Low biodiversity was reflected by both the low number of species and the fact that a few species represent the vast majority in numbers. It is suggested that the area deserves future biological studies, especially in autumn, and a special protection management.     

Demographic parameters of black-browed albatrosses Thalassarche melanophris from the Falkland Islands

Black-browed albatrosses Thalassarche melanophris are currently classified as globally endangered. The most important populations of this species are believed to be declining due to, amongst other factors, unsustainable levels of incidental mortality in fishing gear. However, detailed demographic data are lacking for several critical populations, including the largest of all, nesting in the Falkland Islands. Here, we present data from the first Falkland Islands detailed demographic study (at New Island) and show that, from 2003 to 2009, the mean adult survival probability was 0.942 (95% CI: 0.930–0.952). Nesting frequency of adults is amongst the highest recorded for Thalassarche albatrosses and breeding success (0.564 chicks per egg) is within normal values. The nesting population in the intensively studied plots experienced an increase of 4% per year from 2004 to 2009. These results indicate that the Falklands population may not be as threatened as previously supposed, although studies from more sites and a longer time series are needed to confirm or refute this. The high survival rates may partly reflect recent efforts to mitigate bycatch made by the Falkland Islands and other fisheries in the region. The reinforcement of such initiatives may be critical to buffer the black-browed albatross population against ecosystem shifts and natural disasters (such as harmful algal blooms) that will likely become more frequent with ongoing global changes.     

The breeding biology of the white-chinned petrel Procellaria aequinoctialis at South Georgia

Aspects of the breeding biology of the whitc-chinned petrel Procellaria aequinoctialis were studied at South Georgia. Laying, hatching and fledging dates, together with egg and adult measurements, rate of weight loss of eggs and incubating adults and data on the duration of incubation shifts were recorded. Chick growth in weight, tarsus length, culmen length and wing length from hatching to fledging was followed in detail and breeding success and causes of mortality documented. Comparisons are made with other populations of the species, other members of the genus Procellaria and other Procellariiformes. Despite being the largest burrow-nesting petrel in the world, P. aequinoctialis conforms well to the overall relationships between body weight and breeding parameters for Procellariiformes in general.     

Spatial and temporal distribution of Antarctic fur seals (Arctocephalus gazella) on the breeding grounds at Bird Island,South Georgia

Summary The number of Antarctic fur seals (Arctocephalus gazella) in the area of the Scotia Sea has been increasing. Observation of their distribution on the breeding grounds are important to help design and interpret censuses. These seals are highly polygynous. In the study area at Bird Island, South Georgia, females give birth and are mated in densely packed colonies located along the shore close to the tideline. Males establish territories in this area, but are also found further inland. This study examined the density-dependent processes regulating the instantaneous size of the male breeding population; the distribution of males in relation to space and the number of females available to be mated and the effect of gregarious behaviour of females on male dispersion. Males were only territorial on the beaches in the areas where most females gave birth and subsequently had their post-partum oestrus. There was an apparent lower (19–20 m²) and upper (40 m²) limit to territory size. Males were excluded from the beach areas when the average density on the beaches was greater than 5 males per 200 m². An asymptotic density of 10–11 males per 200 m² was reached on the beaches and 4–5 males per 200 m² elsewhere. These two asymptotic densities may represent the upper and lower limits of density for a territorial system of dispersion. A model of the temporal changes in female numbers suggested that the total number of females occupying an area of beach during the mating period was approximately twice the number at the peak of the season. There was no indication that males compensated for low female density by increasing territory size. Females and pups became more dispersed after the end of the mating period. It is suggested that one function of gregariousness in females is as a mechanism for mate selection. This study also has implications for methods used to measure population size.     

Flying at no mechanical energy cost: disclosing the secret of wandering albatrosses

ALBATROSSES DO SOMETHING THAT NO OTHER BIRDS ARE ABLE TO DO: fly thousands of kilometres at no mechanical cost. This is possible because they use dynamic soaring, a flight mode that enables them to gain the energy required for flying from wind. Until now, the physical mechanisms of the energy gain in terms of the energy transfer from the wind to the bird were mostly unknown. Here we show that the energy gain is achieved by a dynamic flight manoeuvre consisting of a continually repeated up-down curve with optimal adjustment to the wind. We determined the energy obtained from the wind by analysing the measured trajectories of free flying birds using a new GPS-signal tracking method yielding a high precision. Our results reveal an evolutionary adaptation to an extreme environment, and may support recent biologically inspired research on robotic aircraft that might utilize albatrosses' flight technique for engineless propulsion.     

Field metabolic rates of black-browed albatrosses Thalassarche melanophrys during the incubation stage

Field metabolic rates (FMR) and activity patterns of black-browed albatrosses Thalassarche melanophrys were measured while at sea and on nest during the incubation stage at Kerguelen Island, southwestern Indian Ocean. Activity-specific metabolic rates of five albatrosses at sea (FMR_at-sea) were measured using doubly labeled water (DLW), and by equipping birds with wet-dry activity data loggers that determined when birds were in flight or on the water. The metabolic rates of four birds incubating their eggs (FMR_on-nest) were also measured using DLW. The mean±SD FMR_at-sea of albatrosses was 611±96 kJ kg⁻¹ d⁻¹ compared to FMR_on-nest of 196±52 kJ kg⁻¹ d⁻¹. While at sea, albatrosses spent 52.9±8.2% (N=3) of their time in flight and they landed on the water 41.2±13.9 times per day. The FMR of black-browed albatrosses appear to be intermediate to that of three other albatross species. Based on at-sea activity, the power requirement of flight was estimated to be 8.7 W kg⁻¹ (or 4.0×predicted BMR), which is high compared to other albatross species, but may be explained by the high activity levels of the birds when at sea. The FMR_at-sea of albatrosses, when scaled with body mass, are lower than other seabirds of similar body size, which probably reflects the economical nature of their soaring flight.     

Annual variation in breeding biology of macaroni penguins, Eudyptes chvysolophus, at Bird Island, South Georgia

The breeding biology of the macaroni penguin, Eudyptes chrysolophus , was studied over four years, 1976 and 1986–88, at Bird Island, South Georgia. Birds were migratory, being absent during winter (May to September). Arrival at the colony was highly synchronous between years: 14–23 October, over a 7-year period. The pre-breeding, incubation and chick-brooding period was characterized by long fasts ashore, for 36 and 39 days in males and 41 days in females, alternating with long periods at sea. Within years egg-laying was highly synchronous: 95% of clutches initiated within 4–6 days. Arrival date and mean egg-laying date were later (by 3 days), and breeding population size lower (by 20%) in 1987, compared to other years. The incubation period was 35 days and comprised three long shifts, the first shared by the male and female, the second by the female and the third by the male. In 1986 and 1988 these were of 12, 12 and 9 days' duration, but in 1987 the first shift was significantly shorter: 9 days. Chicks creched at 23–25 days of age and fledged at 60 days of age. Neither chick age nor weight at creching or fledging varied between the years 1986–88. The breeding biology of macaroni penguins at Bird Island is compared with that of other Eudyptes penguins, and with the sympatric gentoo penguin, Pygoscelispupuu. There is little variation in breeding biology within the genus Eudyptes , except in the length of time spent at sea prior to the annual moult. This is much shorter at Bird Island, probably reflecting a greater food availability compared to other localities. Inter-annual variation in certain breeding parameters, e.g. laying date, breeding population size, is much greater in the gentoo penguin than in the macaroni penguin. The shorter breeding season, rearing of only one chick and proportionately lower chick fledging weight in macaroni penguins, may be linked to this species' migratory strategy.     

Annual variation in breeding biology of gentoo penguins, Pygoscelis papua, at Bird Island, South Georgia

The breeding biology of the gentoo penguin, Pygoscelis papua , was studied over a three-year period (1986–1988) at Bird Island, South Georgia, with particular reference to birds of known age or breeding experience. Laying date varied significantly between all three years, being three weeks later in 1987, when the breeding population decreased markedly. Factors involved in the timing of breeding are discussed. Within years egg-laying was highly synchronous: 95% of clutches were initiated in 14·5 days or less. The incubation period was 35 days and the laying interval, between the two eggs, 3·3–3·4 days. Chicks creched when 25–30 days old, and this varied between years, possibly related to food supply and chick growth. Chicks left the colony for the first time between 75 and 85 days of age. The breeding population at Bird Island decreased by 20% and increased by 84% in successive years during the study period. Breeding success (chicks fledged per egg laid) varied between 0·33 and 0·65 within colonies, but for the whole island was very consistent over the three years: 0·45, 0·51 and 0·47. Overall, colony differences were not correlated between years. Disturbance from Antarctic fur seals, Arctocephalus gazella , is suggested as the cause of consistently lower breeding success at one colony. Mean egg weight varied annually, and with age of the breeding bird, nest location and, in one year, with laying date. Young, first-time breeders laid smaller eggs and had lower breeding success compared to older, experienced birds, similar to other seabirds. However, they differed from other species in laying on average earlier than older birds. The relationship between age, egg weight, laying date and breeding success is discussed in relation to predation and seasonal food supply.     

The autumn mesozooplankton community at South Georgia: biomass, population structure and vertical distribution

Mesozooplankton were sampled at shelf and oceanic stations close to South Georgia, South Atlantic during austral autumn 2004 with a Longhurst Hardy Plankton Recorder. Onshelf biomass ranged from 2.18 to 5.75 g DM m^?2 (0–200 m) and was dominated by the small euphausiid Thysanöessa spp. At the oceanic stations (10.57–14.71 g DM m^?2, 0–1,000 m) large calanoids, principally Rhincalanus gigas comprised ～47–52% of biomass. Here Calanus simillimus was still active and reproducing in surface waters (0–11.2 eggs fem day^?1) but R. gigas and Calanoides acutus were largely resident in the warm deep water and undergoing their seasonal descent. A comparison with spring and summer data indicated increased abundance and biomass from spring through to summer followed by a decline towards autumn particularly over the shelf. Autumn values in oceanic waters differed little from summer. Mesozooplankton biomass in the surface 200 m of the oceanic stations as a proportion of that found in the top 1,000 m ranged from 63 to 78% of the total in spring and 62–73% in summer, but was only 23–29% of the total in this study, following redistribution down the water column.