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.     

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.     

Factors affecting the number and mortality of seabirds attending trawlers and long-liners in the Kerguelen area

The factors affecting the number and the mortality rates of seabirds attending long-liners and trawlers fishing in the Kerguelen area were studied during four successive seasons (1994–1997), based on observations carried out onboard by dedicated observers. Twenty-four species of seabirds were observed attending fishing vessels, representing an average of 591 birds/census. The total numbers attending varied mainly according to the year, the cloud cover and the presence of offal from long-liners. The dumping of offal increased the numbers of birds attending the vessel, especially when the offal could be easily handled by birds. The activity of the vessels also affected the numbers attending, birds being more abundant during line setting and trawl hauling. White-chinned petrels were the most abundant ship-following seabirds, followed by black-browed albatrosses, giant petrels and cape petrels. The number of white-chinned petrels, black-browed and grey-headed albatrosses attending fishing vessels increased in the time between spring and autumn, whereas it was the reverse situation for giant petrels and cape petrels. Four species of seabirds were caught by fishing gear, mainly by long-lines: white-chinned petrels, and black-browed, grey-headed and wandering albatrosses. Taking into account the number of birds from each species attending long-liners and known to be potential by-catch, some species appear to be more susceptible to being caught than others. White-chinned and grey-headed albatrosses are caught in much higher proportions than the numbers present, whereas black-browed albatrosses are caught in lower numbers. Giant petrels are abundant around long-liners but were never caught. In long-liners, most birds were killed when the lines were set during the day or when the deployment of the scaring device was not successful, with an overall figure of 0.47 birds/1000 hooks. Only one albatross was caught when the lines were set during the night. White-chinned petrels represented 92.2% of all birds killed by long-liners. The number of birds caught varied significantly among months and among years. The type of bait used also affected the catch rate. The catch rate was related to the number of birds attending the long-liner only for black-browed albatrosses. Most birds killed by trawlers were entangled by the netsonde cable. The efficiency of mitigation measures in order to reduce seabird mortality is discussed and it is stressed that night setting is the most efficient way to reduce mortality and should be enforced everywhere when possible. However, further methods should be developed to reduce the mortality of species active at night, especially white-chinned petrels whose populations in the Indian Ocean may by threatened by long-line fisheries. Accepted: 15 October 1999     

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.     

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     

Copulation behaviour and paternity in shy albatrosses (Thalassarche cauta)

We investigated the mating system of shy albatrosses Thalassarche cauta by combining behavioural observations during the pre-laying period with genetic paternity analysis. Genetic data on the mating systems of several procellariiform seabirds have recently become available, but data on the reproductive behaviour of these species are rarely obtained. Our main aims were to describe the copulatory behaviour of this species and identify how males achieve within-pair and extra-pair paternity (EPP). Most copulations occurred on the nest, were unforced and were within-pair. Females controlled the success of copulations and were observed soliciting extra-pair matings. Within-pair and extra-pair copulations were behaviourally similar. A low frequency (7–10%, n =29 chicks) of EPP was detected despite male use of frequent copulation as a paternity guard. The pre-laying foraging exodus of female shy albatrosses differed from that in other albatrosses: it was relatively short in length, lasting c . 2 days, and within-pair copulations occurred after the female's return 2 days before laying. This may reflect the close proximity of feeding grounds to the breeding colony.     

The influence of synoptic weather patterns on the at-sea behaviour of three species of albatross

The relationships between behaviour and synoptic-scale weather patterns and wind speeds for wandering (Diomedea exulans), black-browed (D. melanophrys) and light-mantled sooty (Phoebetria palpebrata) albatrosses were examined using observations collected from ships in the Southern Ocean south of Tasmania, Australia. Wandering albatrosses were found in high-pressure systems in significantly higher numbers than expected than in low-pressure systems. Wandering albatrosses were also found more frequently at lower latitudes than were black-browed and light-mantled sooty albatrosses. Light-mantled sooty and black-browed albatrosses showed no significant association with high- or low-pressure systems, suggesting that they can use either pressure system for travel. High-pressure systems may act as "traps" for wandering albatrosses, due to the lower wind speeds generally associated with these systems.     

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     

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.     

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.     

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.     

Breeding synchrony and extrapair paternity in a species with alternative reproductive strategies

Breeding synchrony may affect the tradeoff between pursuing multiple mates and avoiding paternity loss, translating into differences in the rate of extrapair paternity (EPP). However, diverse empirical relationships between breeding synchrony and EPP remain challenging to explain. We examined whether the relationship between breeding synchrony and EPP varied with male morph, age, body size, or breeding density in the white‐throated sparrow Zonotrichia albicollis. In this species, males of two genetically determined morphs pursue alternative mating strategies. Breeding synchrony positively correlated with EPP within polygamous white morph males, which have high rates of EPP and cuckoldry, but was unrelated to EPP within tan morph males, which prioritize mate guarding and paternal care. As previously reported, males that gained EPP were primarily white males. Males gained EPP more often than expected by chance during their mate's fertile period and on neighboring territories. Since extrapair copulation appears primarily male‐driven in this species, these results indicate that white males focus extra‐pair mating effort during periods of high synchrony and during their mates’ fertile periods, even at the expense of paternity loss within their own nests. Breeding density, male age, and male size did not modify the relationship between breeding synchrony and EPP. However, older white males had higher cuckoldry rates, perhaps reflecting declines in performance associated with senescence. Results suggest that, even within species, mating strategy may modify how breeding synchrony affects rates of EPP, with positive relationships manifest only within subsets of individuals that pursue a strategy of polygyny at the expense of paternity loss.     

Giant petrels as predators of albatross chicks

Giant petrels Macronectes spp. are not thought to be important predators of albatross chicks, although they are known to kill pre-fledging Thalassarche and Phoebetria albatrosses. We report the first records of predation of healthy great albatross Diomedea spp. chicks, killing wandering albatrosses D. exulans at night on sub-Antarctic Marion Island. Breeding success of this species has decreased markedly in the area where attacks occurred, suggesting that giant petrel predation events are a recent phenomenon. Mouse attacks on wandering albatross chicks may have contributed to the development of this hunting technique. We also report the first observations of giant petrel predation on pre-fledging grey-headed albatross T. chrysostoma chicks as well as additional records of sooty albatross P. fusca chicks being targeted. Only adult northern giant petrels M. halli have been confirmed to kill albatross chicks on Marion Island. Given the threatened status of wandering albatrosses, and the importance of Marion Island for this species, monitoring of their breeding success is necessary to assess whether the predation of chicks by giant petrels spreads around the island.     

Extrapair paternity in a flock-living passerine, the vinous-throated parrotbill Paradoxornis webbianus

Understanding the factors that affect the occurrence of extrapair paternity (EPP) is one of the central issues in sexual selection. We investigated genetic parentage and the ecological factors affecting patterns of EPP in the vinous-throated parrotbill, Paradoxornis webbianus , a flock-living species with double broods. Using microsatellite DNA fingerprinting, we determined parentage of 246 offspring in 50 broods over two years (2005 and 2006). Nineteen offspring (8%) from 13 broods (26%) were sired by extrapair males and one offspring (0.4%) was probably the result of intraspecific brood parasitism. The prevalence of EPP varied significantly through the breeding season: 95% of broods with EPP (12/13) occurred in the first of two laying peaks. Parentage assignment revealed that half of extrapair males (6/12) were adjacent neighbours. The distribution of EPP was not significantly related to the ecological factors including breeding density and breeding synchrony. Instead, we suggest that social characteristics such as flocking and weak territoriality may determine the observed pattern of EPP in this study.     

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.     

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.     

Origin, age, sex and breeding status of wandering albatrosses (Diomedea exulans), northern (Macronectes halli) and southern giant petrels (Macronectes giganteus) attending demersal longliners in Falkland Islands and Scotia Ridge waters, 2001–2005

A total of 547 sightings of 291 banded wandering albatrosses Diomedea exulans and 21 sightings of 14 banded giant petrels Macronectes spp. were made from toothfish longliners operating on the southern Patagonian Shelf during 2001–2005. This included 25% of the wandering albatrosses with Darvic bands that bred at Bird Island (South Georgia) during this period. Thirteen of the northern Macronectes halli and southern giant petrels Macronectes giganteus had been banded at South Georgia, and there was one sighting of a southern giant petrel from Argentina. Male and female wandering albatrosses of all age classes except young birds (<15 years old) were equally likely to attend longline vessels. Most sightings of all age classes were made during the incubation period and fewest during the brood period. Eighty-six percent of birds sighted had bred at least once before, with half currently breeding and half on sabbatical (i.e. between breeding attempts). Almost half of the wandering albatrosses were sighted on more than one occasion. The data confirms that the southern Patagonian shelf is an important foraging area for wandering albatrosses and northern and southern giant petrels, and that some individuals show consistent associations in multiple years with longline vessels fishing in the region.     

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.     

Surviving with low genetic diversity: the case of albatrosses

Low genetic diversity is predicted to negatively impact species viability and has been a central concern for conservation. In contrast, the possibility that some species may thrive in spite of a relatively poor diversity has received little attention. The wandering and Amsterdam albatrosses (Diomedea exulans and Diomedea amsterdamensis) are long-lived seabirds standing at an extreme along the gradient of life strategies, having traits that may favour inbreeding and low genetic diversity. Divergence time of the two species is estimated at 0.84 Myr ago from cytochrome b data. We tested the hypothesis that both albatrosses inherited poor genetic diversity from their common ancestor. Within the wandering albatross, per cent polymorphic loci and expected heterozygosity at amplified fragment length polymorphisms were approximately one-third of the minimal values reported in other vertebrates. Genetic diversity in the Amsterdam albatross, which is recovering from a severe bottleneck, was about twice as low as in the wandering albatross. Simulations supported the hypothesis that genetic diversity in albatrosses was already depleted prior to their divergence. Given the generally high breeding success of these species, it is likely that they are not suffering much from their impoverished diversity. Whether albatrosses are unique in this regard is unknown, but they appear to challenge the classical view about the negative consequences of genetic depletion on species survival.     

Chick metabolic rate and growth in three species of albatross: a comparative study

The relative importance of genetic vs. environmental factors in determining the pattern of avian post-embryonic development is much debated. Previous cross-fostering of albatrosses suggested that although inter-specific variation in growth rate was determined primarily by differences in dietary energy content, species-specific constraints might have evolved that could limit maximal growth, even in chicks fed at similar rates and on similar diets. This study aimed to determine whether intrinsic differences in resting metabolic rate were apparent during the linear phase of growth in chicks of three species (black-browed, grey-headed and light-mantled sooty albatrosses). There was a gradual increase in absolute, and a reduction in mass-specific metabolic rate from 5.0 W kg(-1) during the earliest part of linear growth, to 3.5 W kg(-1) by the time chicks reached peak mass. These values are considerably higher than in resting adults of comparable or lower mass, presumably reflecting the large size and high metabolic demand of organs involved in rapid nutrient processing and tissue synthesis by chicks. The lack of any detectable inter-specific variation in the pattern of metabolic rate changes casts some doubt on the existence of fundamental differences in growth rate that cannot be attributed simply to differences in dietary energy or nutrient delivery rate.