The food and feeding ecology of the giant petrels Macronectes halli and M. giganteus at South Georgia

Giant petrels are the chief scavenging seabirds in the Southern Ocean. Quantitatively analysed regurgitations from chicks of both Macronectes halli and M. giganteus at Bird Island, South Georgia throughout the chick rearing period consisted predominantly of adult Macaroni penguins Eudyptes chrysolophus. Significant quantities of burrowing petrels and krill were taken but the amount varied in different years. Only male M. halli fed regularly at fur seal carcases. Although 10 species were found in regurgitations, squid were unimportant in the diet. Fish are possibly more important during the winter.
Intersexual differences in diet were of greater significance than interspecific ones with males taking more carrion and females more crustaceans. Chicks were fed during both day and night and in both species males received significantly more food than females during the fledging period. M. giganteus chicks received more than M. halli at all ages.
Despite their extensive reliance on carrion both species take a variety of prey and this may be especially important during the winter when penguin and seal carrion is scarce.     

Sexual size dimorphism in northern giant petrels: ecological correlates and scaling

Northern giant petrels ( Macronectes halli ) are among the largest and most sexually size dimorphic species of seabirds, with females being only 80% the mass of males. Both sexes scavenge on seal and penguin carrion in the sub-Antarctic ecosystem, but during the breeding season females also feed extensively on other marine food resources and show more pelagic habits than males. The outstanding sexual segregation in foraging and feeding ecology in northern giant petrels suggests that mechanisms maintaining sexual size dimorphism by ecological factors may be operating. I evaluated this possibility by examining ecological correlates with body size and by static allometry analyses. Fledging sex ratio in four consecutive years did not depart from parity. There was no assortative mating by size neither association between the male size with the breeding performance. By contrast, smaller females raised their chick in better condition. Moreover, bill size showed a size dimorphism beyond that expected by body size dimorphism, i.e. when controlling for body mass, males showed relatively longer bill than females. This trait did not deviate from isometry with respect to body size and its phenotypic variability was low, suggesting that the disproportionately large bill of males is related to their more scavenging life style compared to females. In general, the increase and maintenance of sexual size dimorphism in giant petrels is more consistent with an ecological causation rather than a result of sexual selection.     

Species and sexual isolating mechanisms in sibling species of giant petrels Macronectes

Summary The two sibling species of giant petrels Macronectes halli and M. giganteus are the dominant avian scavengers in the Southern Ocean ecosystem. They breed sympatrically at a number of sub-Antarctic sites. This paper synthesises data from a detailed study at South Georgia to examine the importance of various interspecific and intersexual differences between these closely related species. Morphological, breeding, dietary, feeding and moult characteristics were investigated. The most significant interspecific differences are a six week separation in the onset of breeding and the importance of seal carrion to male M. halli, the earlier breeding species. There is a strong sexual size dimorphism with females being only 75–80% of the weight of males in both species. Intersexual dietary differences are stronger than interspecific ones and females also have later primary moult schedules. Reproductive isolation and ecological adaptations are discussed in relation to the present distribution of giant petrels, comprising a more restricted sub-Antarctic species (M. halli) and a widespread Antarctic species (M. giganteus). It is suggested that the marked sexual dimorphism evolved before the two taxa became specifically distinct.     

Segregation in space and time explains the coexistence of two sympatric sub‐Antarctic petrels

Biological communities are shaped by competition between and within species. Competition is often reduced by inter‐ and intraspecific specialization on resources, such as differencet foraging areas or time, allowing similar species to coexist and potentially contributing to reproductive isolation. Here, we examine the simultaneous role of temporal and spatial foraging segregation within and between two sympatric sister species of seabirds, Northern Macronectes halli and Southern Macronectes giganteus Giant Petrels. These species show marked sexual size dimorphism and allochrony (with earlier breeding by Northern Giant Petrels) but this is the first study to test for differences in foraging behaviours and areas across the entire breeding season both between the two species and between the sexes. We tracked males and females of both species in all breeding stages at Bird Island, South Georgia, to test how foraging distribution, behaviour and habitat use vary between and within species in biological time (incubation, brood‐guard or post‐brood stages) and in absolute time (calendar date). Within each breeding stage, both species took trips of comparable duration to similar areas, but due to breeding allochrony they segregated temporally. Northern Giant Petrels had a somewhat smaller foraging range than Southern Giant Petrels, reflecting their greater exploitation of local carrion and probably contributing to their recent higher population growth. Within species, segregation was spatial, with females generally taking longer, more pelagic trips than males, although both sexes of both species showed unexpectedly plastic foraging behaviour. There was little evidence of interspecific differences in habitat use. Thus, in giant petrels, temporal segregation reduces interspecific competition and sexual segregation reduces intraspecific competition. These results demonstrate how both specialization and dynamic changes in foraging strategies at different scales underpin resource division within a community.     

Hybridization and Back-Crossing in Giant Petrels (Macronectes giganteus and M. halli) at Bird Island,South Georgia,and a Summary of Hybridization in Seabirds

Hybridization in natural populations provides an opportunity to study the evolutionary processes that shape divergence and genetic isolation of species. The emergence of pre-mating barriers is often the precursor to complete reproductive isolation. However, in recently diverged species, pre-mating barriers may be incomplete, leading to hybridization between seemingly distinct taxa. Here we report results of a long-term study at Bird Island, South Georgia, of the extent of hybridization, mate fidelity, timing of breeding and breeding success in mixed and conspecific pairs of the sibling species, Macronectes halli (northern giant petrel) and M. giganteus (southern giant petrel). The proportion of mixed-species pairs varied annually from 0.4–2.4% (mean of 1.5%), and showed no linear trend with time. Mean laying date in mixed-species pairs tended to be later than in northern giant petrel, and always earlier than in southern giant petrel pairs, and their breeding success (15.6%) was lower than that of conspecific pairs. By comparison, mixed-species pairs at both Marion and Macquarie islands always failed before hatching. Histories of birds in mixed-species pairs at Bird Island were variable; some bred previously or subsequently with a conspecific partner, others subsequently with a different allospecific partner, and some mixed-species pairs remained together for multiple seasons. We also report the first verified back-crossing of a hybrid giant petrel with a female northern giant petrel. We discuss the potential causes and evolutionary consequences of hybridization and back-crossing in giant petrels and summarize the incidence of back-crossing in other seabird species.     

Sexual dimorphism,moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97

Ryan, P.G. 1999. Sexual dimorphism, moult and body condition of seabirds killed by longline vessels around the Prince Edward Islands, 1996–97. Ostrich 70 (3&4): 187–192.

A total of 393 seabirds from nine species killed by longline fishing in South African waters around the Prince Edward Islands during summer 1996–97 were sexed, measured, and their body fat levels and primary moults scored. Males of Whitechinned Petrels Procellaria aequinoctialis and Greyheaded and (Indian Ocean) Yellownosed Mollymawks Thalassarche chrysostoma and T. [chlororhynchosj bassi averaged 1–9% larger than females. Measures of bill depth were the most dimorphic characters in all three species. I recommend that if only a single measure is taken to estimate sex of adults, bill depth at the base be the standard measure for petrels, and bill depth at the nail be the standard measure for mollymawks. Discriminant functions are described for predicting the sex of unknown individuals; the proportion of misclassified birds ranged from 2–10%. With the exception of giant petrels Macronectes spp., only two young birds were actively moulting their primary flight feathers. Analysis of wear patterns among adult Greyheaded Mollymawk primaries conforms with studies of moult in this species at South Georgia. Fat scores did not differ between sexes. They tended to decrease during the breeding season, but trends were masked by great inter-individual variation. There was no significant relationship between fat scores and the presence of ingested plastics. Plastic debris was found in the stomachs of 90 birds from five species. Incidence in Whitechinned Petrel stomachs (37%) was less than that reported from birds collected off South Africa in the early 1980s (57%), but this is probably due to the predominance of breeding adults. The frequency and size of plastic loads decreased through the breeding season, which is consistent with the hypothesis that inter-generational transfer is important in the dynamics of ingested debris.     

The impacts of introduced House Mice on the breeding success of nesting seabirds on Gough Island

Invasive species are the main threat to island biodiversity; seabirds are particularly vulnerable and are one of the most threatened groups of birds. Gough Island, a UNESCO World Heritage Site in the South Atlantic Ocean, is an Important Bird and Biodiversity Area, and one of the most important seabird colonies globally. Invasive House Mice Mus musculus depredate eggs and chicks of most seabird species on the island, but the extent of their impact has not been quantified. We used field data and bootstrapped normal distributions to estimate breeding success and the number of surviving chicks for 10 seabird species on Gough Island, and compared estimates with those of analogous species from predator‐free islands. We examined the effects of season and nest‐site location on the breeding success of populations on Gough Island, predicting that the breeding success of Gough birds would be lower than that of analogues, particularly among small burrow‐nesting species. We also predicted that winter‐breeding species would exhibit lower breeding success than summer‐breeding species, because mice have fewer alternative food sources in winter; and below‐ground nesters would have lower breeding success than surface nesters, as below‐ground species are smaller so their chicks are easier prey for mice. We did indeed find that seabirds on Gough Island had low breeding success compared with analogues, losing an estimated 1 739 000 (1 467 000–2 116 000) eggs/chicks annually. Seven of the 10 focal species on Gough Island had particularly high chick mortality and may have been subject to intense mouse predation. Below‐ground and winter breeders had lower breeding success than surface‐ and summer‐breeders. MacGillivray's Prion Pachyptila macgillivrayi, Atlantic Petrel Pterodroma incerta and Tristan Albatross Diomedea dabbenena are endemic or near‐endemic to Gough Island and are likely to be driven to extinction if invasive mice are not removed.     

Who’s the boss? Giant petrel arrival times and interspecific interactions at a seal carcass at sub-Antarctic Marion Island

Most northern giant petrels Macronectes halli arrived before southern giant petrels Macronectes giganteus at a seal carcass at sub-Antarctic Marion Island and left earlier. In interspecific interactions, southern giant petrels initiated and dominated encounters. Level of aggression may be related to the degree of satiation. Dominance by southern giant petrels at seal carcasses may help explain why northern giant petrels commence breeding earlier than southern giant petrels in sympatric situations.     

Nocturnal predation of king penguins by giant petrels on the Crozet Islands

Dietary segregation of sympatric seabirds in the Southern Ocean is partly linked to differences in their foraging techniques. We have investigated the activity of giant petrels (Macronectes spp.) in a king penguin (Aptenodytes patagonicus) colony day and night during the austral winter of 2001 on the Crozet Islands. Using an automatic identification system and an infrared video camera, we followed 15 petrels tagged with micro transponders. Our data show that giant petrels predate king penguin chicks during the night. The activity of giant petrels is even slightly higher during nighttime than during the day. In addition, our data show a higher nocturnal activity by northern giant petrels (M. halli) than by southern giant petrels (M. giganteus). These unexpected results raise questions concerning visual adaptations to nocturnal foraging in giant petrels and their potential impact on the sleep, vigilance and crèching behavior of penguin chicks.     

The modern avifauna of the Pitcairn Islands

Excluding the gadfly petrels Pterodroma spp. and the resident landbirds, this paper details the present status of bird species seen during an expedition to the Pitcairn Islands (Ducie, Henderson, Oeno and Pitcairn) in 1991 and early 1992. Ten species were recorded in the Islands for the first time. Several Southern Ocean petrels were recorded, most in the midwinter period of June and July. The populations of Christmas Island shearwaters breeding on Oeno and Ducie laid in synchrony every 9–10 months. The species is one of few with such a sub-annual, synchronized regime. The majority of other seabirds had an annual breeding cycle, laying between May and October. Around one percent of the world population of the bristle-thighed curlew passes the non-breeding season in the Pitcairn Islands.     

The impact of avian predator-scavengers on King Penguin Aptenodytes patagonicus chicks at Marion Island

While ashore King Penguins Aptenodytes patagonicus are fed upon by a guild of five predator-scavenger seabirds. During the winter (April-October) male Southern Giant Petrels Macronectes giganteus killed an estimated 6430 (11.2%) of Marion Island's King Penguin chicks, although most birds only scavenged in the colonies. The rate of predation varied, with peaks in April-May and in September. The proportion of successful attacks was 22.7%. There was a strong correlation between colony size and the rate of accumulation of chick corpses. Kelp Gulls Larus dominicanus and Lesser Sheathbills Chionis minor also scavenged penguin corpses but Northern Giant Petrels M. halli and female Southern Giant Petrels rarely entered the colonies. During the summer predation was mainly by Sub-Antarctic Skuas Catharacta lonnbergi which took eggs and small chicks.     

Widespread gene flow between oceans in a pelagic seabird species complex

Global‐scale gene flow is an important concern in conservation biology as it has the potential to either increase or decrease genetic diversity in species and populations. Although many studies focus on the gene flow between different populations of a single species, the potential for gene flow and introgression between species is understudied, particularly in seabirds. The only well‐studied example of a mixed‐species, hybridizing population of petrels exists on Round Island, in the Indian Ocean. Previous research assumed that Round Island represents a point of secondary contact between Atlantic (Pterodroma arminjoniana) and Pacific species (Pterodroma neglecta and Pterodroma heraldica). This study uses microsatellite genotyping and tracking data to address the possibility of between‐species hybridization occurring outside the Indian Ocean. Dispersal and gene flow spanning three oceans were demonstrated between the species in this complex. Analysis of migration rates estimated using bayesass revealed unidirectional movement of petrels from the Atlantic and Pacific into the Indian Ocean. Conversely, structure analysis revealed gene flow between species of the Atlantic and Pacific oceans, with potential three‐way hybrids occurring outside the Indian Ocean. Additionally, geolocation tracking of Round Island petrels revealed two individuals travelling to the Atlantic and Pacific. These results suggest that interspecific hybrids in Pterodroma petrels are more common than was previously assumed. This study is the first of its kind to investigate gene flow between populations of closely related Procellariiform species on a global scale, demonstrating the need for consideration of widespread migration and hybridization in the conservation of threatened seabirds.     

Hematology,plasma biochemistry,and serosurvey for selected infectious agents in southern giant petrels from Patagonia,Argentina

In conjunction with reproductive and feeding ecology studies on southern giant petrels (SGP, Macronectes giganteus) blood samples were collected for baseline health evaluations. Twenty-five adult SGP from a breeding colony in Chubut, Argentina, were sampled during two consecutive breeding seasons, 1999-2000 (n = 15) and 2000-01 (n = 10). Values for hematology, plasma biochemistry, and minerals are described for 20 birds in apparent good physical condition. A serologic survey of exposure to selected infectious agents was also conducted on all 25 birds sampled. Southern giant petrels were serologically negative for evidence of exposure to infectious laryngotracheitis virus, avian encephalomyelitis virus, avian influenza virus, avian reovirus, infectious bursal disease virus, infectious bronchitis virus, paramyxovirus 1, 2, and 3 virus, Chlamydophila, and Aspergillus. Antibodies to avian adenovirus were found in 14% of SGP during the first sampling season, and 60% in the second year. Additionally, all birds were negative for antibodies to Salmonella pullorum at the first sampling date, but 90% had low titers the following breeding season. This study contributes to understanding the health status of South Atlantic seabirds and to establishment of baseline information for SGP. Long-term monitoring of pelagic predator-scavenger seabirds such as SGP should be established for the surveillance of marine ecosystem health.     

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.     

Stable isotope evidence of diverse species-specific and individual wintering strategies in seabirds

Although there is increasing evidence that climatic variations during the non-breeding season shape population dynamics of seabirds, most aspects of their winter distribution and ecology remain essentially unknown. We used stable isotope signatures in feathers to infer and compare the moulting (wintering) habitat of subantarctic petrels breeding at two distant localities (South Georgia and Kerguelen). Petrels showed species-specific wintering habitat preferences, with a similar pattern of latitudinal segregation for all but one taxon. At both localities, delta13C values indicated that blue petrels (Halobaena caerulea) moult in Antarctic waters, South Georgian diving petrels (Pelecanoides georgicus) in the vicinity of the archipelagos and/or in the Polar Frontal Zone and Antarctic prions (Pachyptila desolata) in warmer waters. In contrast, common diving petrels (Pelecanoides urinatrix) showed divergent strategies, with low and high intrapopulation variation at South Georgia and Kerguelen, respectively. Birds from Kerguelen dispersed over a much wider range of habitats, from coastal to oceanic waters and from Antarctica to the subtropics, whereas those from South Georgia wintered mainly in waters around the archipelago. This study is the first to show such striking between-population heterogeneity in individual wintering strategies, which could have important implications for likely demographic responses to environmental perturbation.     

The timing of gonadal development and moult in three raptors with different breeding seasons: effects of gender, age and body condition

Differences between species in breeding seasons are thought to be mediated through differences in their reproductive physiology. Little is known about how the timing and duration of gonadal maturation varies between raptor species, how the timing of moult relates to the gonadal cycle, whether the timing and degree of sexual maturation varies between juveniles and adults or whether body condition has a significant effect. To address these questions, data on gonadal size and moult for adults and juveniles of both sexes of three raptor species were extracted from the Predatory Bird Monitoring Scheme (based on birds found dead by members of the public). The three species, Sparrowhawk Accipiter nisus, Kestrel Falco tinnunculus and Barn Owl Tyto alba, have different ecologies – diurnal bird predator, diurnal mammal predator and nocturnal mammal predator, respectively. All are single‐brooded but have different breeding seasons. The duration of gonadal maturation was markedly different between the species. Barn Owls showed the earliest maturation and the latest gonad regression, and Sparrowhawks the latest maturation and earliest gonad regression. Kestrels were intermediate. In males of all species, the testes remained fully mature throughout their respective breeding seasons. In females, the ovaries remained partially mature throughout the breeding season. Moult started slightly earlier in Sparrowhawks than in Kestrels and coincided with gonadal regression in the two species. Although females of the two species started to moult earlier than males, moult duration was similar between the sexes. Barn Owls showed no distinct annual pattern of moult. In juveniles of all three species, the gonads were smaller than in adults throughout spring and started to mature later. Gonad size in birds that had starved tended to be smaller than in birds dying from other causes, but did not influence the difference in gonad mass between adults and juveniles and between seasons. Body condition had no effect on moult. Whilst ecology has led to the evolution of different breeding seasons, differences between species, and between adults and juveniles, are mediated through adaptive differences in their reproductive physiology.     

Contrasting effects of tropical cyclones on the annual survival of a pelagic seabird in the Indian Ocean

Tropical cyclones are renowned for their destructive nature and are an important feature of marine and coastal tropical ecosystems. Over the last 40 years, their intensity, frequency and tracks have changed, partly in response to ocean warming, and future predictions indicate that these trends are likely to continue with potential consequences for human populations and coastal ecosystems. However, our understanding of how tropical cyclones currently affect marine biodiversity, and pelagic species in particular, is limited. For seabirds, the impacts of cyclones are known to be detrimental at breeding colonies, but impacts on the annual survival of pelagic adults and juveniles remain largely unexplored and no study has simultaneously explored the direct impacts of cyclones on different life‐history stages across the annual life cycle. We used a 20‐year data set on tropical cyclones in the Indian Ocean, tracking data from 122 Round Island petrels and long‐term capture–mark–recapture data to explore the impacts of tropical cyclones on the survival of adult and juvenile (first year) petrels during both the breeding and migration periods. The tracking data showed that juvenile and adult Round Island petrels utilize the three cyclone regions of the Indian Ocean and were potentially exposed to cyclones for a substantial part of their annual cycle. However, only juvenile petrel survival was affected by cyclone activity; negatively by a strong cyclone in the vicinity of the breeding colony and positively by increasing cyclone activity in the Northern Indian Ocean where they spend the majority of their first year at sea. These contrasting effects raise the intriguing prospect that the projected changes in cyclones under current climate change scenarios may have positive as well as the more commonly perceived negative impacts on marine biodiversity.     

NOTES ON THE BIRDS OF THE ISLANDS OF THE BERLENGAS (PORTUGAL), THE DESERTAS AND BAIXO (MADEIRA) AND THE SALVAGES.

The birds seen on the Berlengas, the Desertas, Baixo Island, and the Salvages in June and July 1939 are listed and field notes are given on certain species.
In Puffinis kuhlii borealis incubation was by both sexes. Where it is unmolested by man, as at Grande Salvage, this bird is partly diurnal in visiting the land; where molested, as on the Berlingas' and Desertas, it is nocturnal.
In Bulweria bulwerii incubation is by both sexes, probably in shifts of two or more days.
At Grande Salvage there appears to be a division of the year into specific breeding seasons by which the maximum use is made of the ground–burrows by two if not three species of petrels. This probable breeding competition is discussed, and other examples suggested from the literature of petrels breeding on small tropical and sub–tropical islands.     

Status,trends, and patterns of covariation of breeding seabirds at St Lazaria Island,Southeast Alaska, 1994–2006

Aim We examined patterns of covariation among piscivorous and planktivorous seabirds breeding at St Lazaria Island in order to evaluate their responses to interannual changes in sea surface temperature, a variable that affects marine food webs. In addition, we evaluated seabird population trends for responses to decadal‐scale changes in the marine ecosystem. Location St Lazaria Island, Sitka Sound, Alaska. Methods Established seabird monitoring protocols for the Alaska Maritime National Wildlife Refuge were followed in estimating population trends, the timing of nesting events and the reproductive success of eight species of seabirds between 1994 and 2006. Results  Population increases were noted for storm‐petrels (Oceanodroma furcata and O. leucorhoa), rhinoceros auklets (Cerorhinca monocerata) and glaucous‐winged gulls (Larus glaucescens). We found no population trend for pelagic cormorants (Phalacrocorax pelagicus), but it appeared that populations of common (Uria aalge) and thick‐billed (U. lomvia) murres and of tufted puffins (Fratercula cirrhata) declined. We detected no linear trends in either breeding chronology or reproductive success over the study period for any seabird. All species of piscivorous seabirds apparently responded similarly to environmental cues as there was a positive covariation among species in the timing of nesting. Piscivores tended to nest earlier, and most species had higher rates of reproductive success in years with relatively warm spring sea temperatures. In contrast, planktivorous Leach’s storm‐petrels (O. leucorhoa) tended to nest earlier when spring and summer sea temperatures were relatively cool. Clearly, seabirds at St Lazaria were responding to interannual changes in sea temperatures near the breeding colony, probably as a result of effects on the food webs. Main conclusions Every seabird species we monitored at St Lazaria exhibited significant population trends between 1994 and 2006. For most species there appeared to be a relationship between both the timing of nesting and reproductive rates and spring or summer sea surface temperatures. Responses at both decadal (populations) and interannual (timing and reproductive success) scales make seabirds useful candidates for helping to monitor change in the marine environment.     

Moult in Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma

We recorded the age of individual wing and tail feathers of Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma of known age and breeding status at Bird Island, South Georgia. Breeders and non-breeders of both species moult their rectrices annually. Non-breeders moult primaries biennially. In the first year of a cycle, the outer three and some inner primaries are moulted descendantly; in the next year the inner primaries are moulted ascendantly, starting from primary seven. There is a general progression to moulting equal numbers of primaries in each half of the cycle by the time breeding starts at about 10 years of age. Grey-headed Albatrosses usually moult fewer primaries than Black-browed Albatrosses, particularly as 3-year-olds, when they undertake substantial plumage change in body moult. Most secondaries in Black-browed Albatrosses have been replaced once by age 4 years. Breeding Black-browed Albatrosses continue the moult pattern established as immatures whether they fail or not, as do failed Grey-headed Albatrosses. Successful Grey-headed Albatrosses, which breed again 16 months later, moult their three innermost primaries after breeding in the remainder of the current year and, after a period when moult is interrupted, renew the remaining primaries the following year. Comparisons between species and between failed and successful birds within species indicate that moult rate is not closely linked to the length of the interval between breeding attempts. Interspecies differences are better explained by breeding latitude, with tropical albatrosses moulting twice as fast as sub-Antarctic species, possibly reflecting food availability outside the breeding season.