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.     

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.     

An effective method for trapping scavenging seabirds at sea

ABSTRACT Most studies of seabirds that involve trapping and marking birds are carried out at breeding colonies. This bias toward the breeding period and colony‐based research is partially caused by difficulties in capturing birds at sea. From 2005 to 2007, we used a cast net thrown by hand from a fishing boat to capture albatrosses and petrels at sea in the southwestern Atlantic Ocean. About 500 birds of 13 species were captured, ranging in size from the 30‐g Wilson's Storm‐petrel (Oceanites oceanicus) to the 10‐kg Wandering Albatross (Diomedea exulans). Cast nets are potentially useful for capturing any seabird that can be attracted close to fishing vessels by bait, such as sardines, squid, offal, or shark liver, thrown into the water. Our method was most effective for capturing bold species, such as Cape (Daption capense), Spectacled (Procellaria conspicillata), and White‐chinned (Procellaria aequinoctialis) petrels and Great Shearwaters (Puffinus gravis), but was not effective for capturing shy species, such as Cory's (Calonectris diomedea), Cape Verde (Calonectris edwardsii) and Manx (Puffinus puffinus) shearwaters, species that rarely sit on the water, such as Wilson's Storm‐petrels, Thin‐billed (Pachyptila belcheri) and Antarctic (Pachyptila desolata) prions and Atlantic Petrels (Pterodroma incerta), and species with excellent diving abilities, such as Sooty Shearwaters (Puffinus griseus). For many species of seabirds, cast nets would likely be more efficient for capturing large numbers of birds than other methods, such as hoop nets.     

Estimates of population size of white-chinned petrels and grey petrels at Kerguelen Islands and sensitivity to fisheries

White-chinned petrels Procellaria aequinoctialis and grey petrels Procellaria cinerea are among the most frequently killed seabird species by accidental bycatch, and both species have received strong conservation concern. Data on population size are required to evaluate the impact of bycatch and to establish management plans. We estimated the population size of both species at Kerguelen, Southern Indian Ocean, from 2004 to 2006 by explicitly taking into account detection probability of burrows using distance sampling and burrow occupancy. A total of 31 line-transects were distributed across the eastern part of Kerguelen, representing a total length of 566 km. Detectability was low (from 0.19 to 0.54 for white-chinned petrels, 0.58 for grey petrels). Burrow densities varied from 1.37±0.67 to 25.77±5.23 burrows ha⁻¹ for white-chinned petrels and was 2.78±0.79 burrows ha⁻¹ for grey petrels. For white-chinned petrels, these densities were extrapolated to the entire surface area of vegetation and there were 234 000 (186 000–297 000) active burrows on Kerguelen. For grey petrels, the number of active burrows for the eastern part of Kerguelen was 3400 (1900–5600). Based on these estimates, the potential biological removal method suggests that the additional mortality on birds caused by the fisheries operating around Kerguelen can be considered a serious threat for the species at least at the regional scale of the Southern Indian Ocean, especially for grey petrels.     

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     

The behaviour of seabirds foraging at fishing boats around Shetland

Among the different types of fishing vessels around Shetland, whitefish trawlers attract the largest numbers of scavenging seabirds and provide the most food. Offal was almost all consumed by seabirds, predominantly by Fulmars Fulmarus glacialis , which excluded other species by their aggression. Fulmars generally ignored discarded whole fish, which were mainly taken by Great Black-backed Gulls Larus marinus , Gannets Sula bassana and Great Skuas Catharacta skua . Although flatfish were usually ignored because seabirds found them difficult to swallow and they sank faster, most discarded roundfish were consumed. Herring Gulls L. argentatus , Lesser Black-backed Gulls L. fuscus and Kittiwakes Rissa tridactyla were rarely able to obtain offal or discards. Herring Gulls and Lesser Black-backed Gulls spent much time on the periphery of feeding flocks while Kittiwakes rarely attempted even to join these. Most of the birds at trawlers were in adult plumage, and it is suggested that the low proportion of immature birds present was a further reflection of the highly competitive feeding conditions at trawlers. We suggest that likely changes in fishing practice and seabird population sizes in the immediate future may result in Herring Gulls, Lesser Black-backed Gulls and Great Skuas finding feeding on waste around trawlers increasingly difficult, so they may be further displaced by Fulmars, Gannets and Great Black-backed Gulls.     

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     

Status and distribution of wandering, black-browed and grey-headed albatrosses breeding at South Georgia

Long-term studies at Bird Island, South Georgia, show that numbers of wandering, black-browed and grey-headed albatrosses have been decreasing since the late 1970s. To determine the status of the total South Georgia population, all known colonies were censused in 2003/2004 using a combination of yacht-based digital photography and ground counts. The breeding population estimates from this census are 1,553 pairs of wandering albatross, 74,296 pairs of black-browed albatross and 47,674 pairs of grey-headed albatross. A 30% decline since 1984 was recorded for wandering albatross, and comparison of a sample of black-browed and grey-headed albatross colonies on the mainland of South Georgia photo-censused in both 1985/1986 and 2003/2004 indicates similarly substantial population declines. Unless these decreases can be halted or reversed, doubt will exist as to the long-term viability of these species of albatross at South Georgia.     

Seabird Bycatch in Pelagic Longline Fisheries Is Grossly Underestimated when Using Only Haul Data

Hundreds of thousands of seabirds are killed each year as bycatch in longline fisheries. Seabirds are predominantly caught during line setting but bycatch is generally recorded during line hauling, many hours after birds are caught. Bird loss during this interval may lead to inaccurate bycatch information. In this 15 year study, seabird bycatch was recorded during both line setting and line hauling from four fishing regions: Indian Ocean, Southern Ocean, Coral Sea and central Pacific Ocean. Over 43,000 albatrosses, petrels and skuas representing over 25 species were counted during line setting of which almost 6,000 seabirds attempted to take the bait. Bait-taking interactions were placed into one of four categories. (i) The majority (57%) of bait-taking attempts were “unsuccessful” involving seabirds that did not take the bait nor get caught or hooked. (ii) One-third of attempts were “successful” with seabirds removing the bait while not getting caught. (iii) One-hundred and seventy-six seabirds (3% of attempts) were observed being “caught” during line setting, with three albatross species – Laysan (Phoebastria immutabilis), black-footed (P. nigripes) and black-browed (Thalassarche melanophrys)– dominating this category. However, of these, only 85 (48%) seabird carcasses were retrieved during line hauling. Most caught seabirds were hooked through the bill. (iv) The remainder of seabird-bait interactions (7%) was not clearly observed, but likely involved more “caught” seabirds. Bait taking attempts and percentage outcome (e.g. successful, caught) varied between seabird species and was not always related to species abundance around fishing vessels. Using only haul data to calculate seabird bycatch grossly underestimates actual bycatch levels, with the level of seabird bycatch from pelagic longline fishing possibly double what was previously thought.     

Multivariate effects on seabird bycatch in the legal Patagonian toothfish longline fishery around Crozet and Kerguelen Islands

The effects of temporal, spatial, environmental and operational effects on seabird incidental mortality in the legal Patagonian toothfish longline fishery operating, between 2003 and 2006, in French exclusive economic zones of Crozet and Kerguelen Islands were analysed. During the study period, the mean bycatch rate varied from 0.05 to 0.12 birds per 1,000 hooks. Two species were concerned by incidental mortality: white-chinned petrels (88%) and grey petrels (11.5%). Males of white-chinned petrel seemed more at a risk than females. Logbooks data tended to underreport mortality when compared with dedicated fishery observers. The results indicate that temporal (season or phenology) and spatial (area) factors reflecting mortality risk for seabirds played the most significant role in the incidental mortality of the two species. Operational (integrated weight mainline, number of scaring lines and number of hooks hauled) and environmental factors (wind/vessel angle, moon brightness) were also influential, although less significantly, in increasing this mortality risk. Our two steps analyses by separately modelling the probability of presence and the abundance given presence suggest that the decrease in seabird bycatch over the period was mainly due to an important decrease in probability (occurrence) of mortality.     

Refining Tori lines to further reduce seabird mortality associated with demersal trawlers in the South Atlantic

Fisheries-related mortality of globally threatened albatrosses and petrels in the Falkland Islands is largely attributed to collisions with warp cables on stern trawlers. Following the introduction Tori lines, also known as bird-scaring lines, in 2004, incidental mortality has been substantially reduced. However, there is still a requirement for further development and optimisation of mitigation methods. In this paper, we present results of sea trials to compare the efficacy of a modified Tori line design with the initial (2004) design. The modified design resulted in 28 and 33% fewer warp contacts than the 2004 design for all species, and high-risk species (black-browed albatross and giant petrels), respectively. Surprisingly, these differences were not found to be statistically significant. However, the modified design was more effective at reducing interactions with the Tori lines themselves. The modified design provided better coverage and protection of the warps, particularly in crosswinds, and was safer for the crew to use. Our study highlights the importance of continued research and development in the field of bycatch mitigation and has relevance for trawl fisheries globally.     

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.     

Albatrosses Following Fishing Vessels: How Badly Hooked Are They on an Easy Meal?

Fisheries have major impacts on seabirds, both by changing food availability and by causing direct mortality of birds during trawling and longline setting. However, little is known about the nature and the spatial-temporal extent of the interactions between individual birds and vessels. By studying a system in which we had fine-scale data on bird movements and activity, and near real-time information on vessel distribution, we provide new insights on the association of a threatened albatross with fisheries. During early chick-rearing, black-browed albatrosses Thalassarche melanophris from two different colonies (separated by only 75 km) showed significant differences in the degree of association with fisheries, despite being nearly equidistant to the Falklands fishing fleet. Most foraging trips from either colony did not bring tracked individuals close to vessels, and proportionally little time and foraging effort was spent near ships. Nevertheless, a few individuals repeatedly visited fishing vessels, which may indicate they specialise on fisheries-linked food sources and so are potentially more vulnerable to bycatch. The evidence suggests that this population has little reliance on fisheries discards at a critical stage of its nesting cycle, and hence measures to limit fisheries waste on the Patagonian shelf that also reduce vessel attractiveness and the risk of incidental mortality, would be of high overall conservation benefit.     

Seasonal sexual segregation in two Thalassarche albatross species: competitive exclusion,reproductive role specialization or foraging niche divergence?

Sexual segregation by micro- or macrohabitat is common in birds, and usually attributed to size-mediated dominance and exclusion of females by larger males, trophic niche divergence or reproductive role specialization. Our study of black-browed albatrosses, Thalassarche melanophrys, and grey-headed albatrosses, T. chrysostoma, revealed an exceptional degree of sexual segregation during incubation, with largely mutually exclusive core foraging ranges for each sex in both species. Spatial segregation was not apparent during brood-guard or post-guard chick rearing, when adults are constrained to feed close to colonies, providing no evidence for dominance-related competitive exclusion at the macrohabitat level. A comprehensive morphometric comparison indicated considerable species and sexual dimorphism in wing area and wing loading that corresponded, both within and between species, to broad-scale habitat preferences relating to wind strength. We suggest that seasonal sexual segregation in these two species is attributable to niche divergence mediated by differences in flight performance. Such sexual segregation may also have implications for conservation in relation to sex-specific overlap with commercial fisheries.     

Predator–prey interactions: why do larger albatrosses eat bigger squid?

The relationship between predator sizes and prey sizes is well documented for terrestrial but rarely for marine ecosystems. We show that wandering albatrosses, the biggest albatross species, feed on larger cephalopod prey than those consumed by smaller albatrosses (grey-headed and black-browed albatrosses). This reflects differences in timing of breeding, foraging ecology and their feeding methods. Wandering albatrosses breed later in the year, during the austral winter, than smaller albatrosses (therefore catching older squid) and forage most of the year in Antarctic open waters, sub-Antarctic, subtropical and tropical waters, overlapping minimally with the smaller albatrosses' foraging range while breeding. Also, wandering albatrosses mostly scavenge whereas smaller albatrosses feed more on live prey. Prey ecology may also play a key role because many squid species might experience post-spawning mortality during the austral winter, becoming easily available to wandering albatrosses. Spawning in winter can be linked to predator avoidance (i.e. reduction in mortality in winter by avoiding pelagic predators) and would allow squid larvae to develop and take advantage of the high productivity (i.e. Antarctic phytoplankton bloom) in spring and at the beginning of summer. Thus, aspects of prey and predator ecology may combine to generate observed differences in prey size.     

Fowl play and the price of petrel: long-living Procellariiformes have peroxidation-resistant membrane composition compared with short-living Galliformes

The membrane pacemaker hypothesis predicts that long-living species will have more peroxidation-resistant membrane lipids than shorter living species. We tested this hypothesis by comparing the fatty acid composition of heart phospholipids from long-living Procellariiformes (petrels and albatrosses) to those of shorter living Galliformes (fowl). The seabirds were obtained from by-catch of commercial fishing operations and the fowl values from published data. The 3.8-fold greater predicted longevity of the seabirds was associated with elevated content of peroxidation-resistant monounsaturates and reduced content of peroxidation-prone polyunsaturates and, consequently, a significantly reduced peroxidation index in heart membrane lipids, compared with fowl. Peroxidation-resistant membrane composition may be an important physiological trait for longevous species.     

Dietary segregation of krill-eating South Georgia seabirds

The diets of six of the main seabird species (two petrels, two albatrosses, two penguins) breeding at Bird Island, South Georgia were studied simultaneously during the chick-rearing period in 1986. For five species, Antarctic krill Euphausia superba was the main food (39–98% by mass); grey-headed albatrosses took mainly the ommastrephid squid Martialia hyadesi (71%) and only 16% krill. The size of the krill taken was similar between seabird species, although there were small but significant differences between penguins and the other species. Sex and reproductive status of krill, however, was different between all seabird species, reflecting some combination of differences in foraging ranges, selectivity by predators, or differences in escape responses of krill. For the krill-eating species, the rest of the diet varied substantially between species, comprising Martialia and nototheniid fish (blackbrowed albatross and, along with lanternfish, white-chinned petrel), lanternfish and amphipods (Antarctic prion and macaroni penguin), and icefish (gentoo penguin). Long-term data on breeding success and information on diet in 5–10 other years suggest that in 1986 seabird diet and reproductive performance was indicative of a year of good availability of krill around South Georgia. In such circumstances, ecological segregation between krill-eating species appears to be maintained chiefly by differences in foraging range and feeding methods, which are reviewed. This situation is rather different from the few studies of seabird communities elsewhere, where prey type and size are believed to be the main mechanisms of dietary segregation.     

Seabirds attending bottom long-line fishing off southeastern Brazil

Flocks of seabirds attending commercial bottom long-line fishing operations on the coastal shelf off southeastern Brazil show a greater species diversity during the summer than the winter (16 v 9 species), although the number of birds per flock tended to be greater during the cold season. During the summer, the Spectacled Petrel Procellaria aequinoctialis conspi-cillata was the commonest species, followed by the Great Shearwater Puffinus gravis, Cory's Shearwater Calonectris diomedea, skuas Stercorarius spp. and the Yellow-nosed Albatross Diomedea chlororhynchus. During the winter, the White-chinned Petrel Procellaria aequin-octialis, Yellow-nosed Albatross and Black-browed Albatross Diomedea melanophrys were jointly the commonest species. Marked differences in the relative abundance of species were observed between the different sampling periods, probably because of migratory movements but also because of seasonal shifts of the sea currents and the influence of cold fronts. Waters off southeastern Brazil are important feeding areas for some seabird populations nesting in the Tristan da Cunha and Gough group, especially for nonbreeding Spectacled Petrels and post-breeding Yellow-nosed Albatrosses.     

Differences in the timing and extent of annual moult of black-browed albatrosses Thalassarche melanophris living in contrasting environments

Moult entails costs related to the acquisition of energy and nutrients necessary for feather synthesis, as well as the impact of reduced flight performance induced by gaps in the wing plumage. Variation in moult strategies within and between populations may convey valuable information on energetic trade-offs and other responses to differing environmental constraints. We studied the moult strategies of two populations of a pelagic seabird, the black-browed albatross Thalassarche melanophris, nesting in contrasting environments. According to conventional wisdom, it is exceptional for albatrosses (Diomedeidae) to moult while breeding. Here we show that black-browed albatrosses breeding on the Falklands regularly moult primaries, tail and body feathers during chick-rearing, and the majority of those at South Georgia show some body feather moult in late chick-rearing. The greater moult-breeding overlap at the Falklands allows the birds to annually renew more primary feathers than their counterparts at South Georgia. The results of the present paper, pooled with other evidence, suggest that black-browed albatrosses from South Georgia face a more challenging environment during reproduction. They also serve to warn against the uncritical acceptance of conventional ideas about moult patterns when using feathers to study the ecology of seabirds and other migrants for which there is scant information at particular stages of the annual cycle.     

Phylogeny of "Philoceanus complex" seabird lice (Phthiraptera: Ischnocera) inferred from mitochondrial DNA sequences

The Philoceanus complex is a large assemblage of lice that parasitise procellariiform seabirds (petrels, albatrosses, and their relatives). We obtained mitochondrial 12S rRNA and cytochrome oxidase I DNA sequences from 39 species from diverse hosts and localities. Resolution of deeper relationships between genera was limited, however there is evidence for two major clades, one hosted by albatrosses, the other by petrels. Based on our results, the genera hosted by albatrosses are excellent candidates for detailed analysis of cospeciation. Our results also suggest that a previous estimate of a 5-fold difference in the relative rate of sequence evolution in lice and their avian hosts is an artefact of limited taxonomic sampling.