Fish prey of the Wandering Albatross Diomedea exulans at South Georgia

Summary The fish diet (45% of total diet by weight) of Wandering Albatrosses rearing chicks at South Georgia during the austral winters of 1983 and 1984 was investigated using otoliths retrieved from regurgitations. These provide the first quantitative data for this species and for any albatross. By number of identified otoliths (32% could be identified only as ?Macrouridae and ?Moridae), Pseudochaenichthys georgianus (35%), Muraenolepis microps (33%) and Chaenocephalus aceratus (20%) predominated, with Notothenia gibberifrons, Pagothenia hansoni and Champsocephalus gunnari (together 12%) also present. Composition by weight (estimated from otolith length) of the main species was Pseudochaenichthys 51%, Muraenolepis 14%, Chaenocephalus 27%; if digestion and wear had reduced otoliths by 10% the values would be Pseudochaenichthys 54%, Chaenocephalus 25%, Muraenolepis 13%. Composition by weight (actual or corrected values) was almost identical between years but epipelagic fish were significantly more abundant in 1983 than 1984. All identified fish eaten by Wandering Albatrosses are common on the South Georgia continental shelf and most of them are caught in the commercial fishery there. However, two of the three main target species of this fishery in 1983–1984, Notothenia rossii and Champsocephalus were not, or rarely, caught by Wandering Albatrosses. It seems unlikely, therefore, that the albatrosses depend greatly on the fishery for acquisition of fish prey but how they catch several species, including Muraenolepis, which are mainly benthic in habit is unknown.     

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.     

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.     

Interannual variability in the feeding of ice fish (Notothenioidei, Channichthyidae) in the southern Scotia Arc and the Antarctic Peninsula region (CCAMLR Subareas 48.1 and 48.2)

We have studied the annual variation in food intake of three sub-Antarctic ice fish species (Champsocephalus gunnari, Chaenocephalus aceratus, and Pseudochaenichthys georgianus) and three high-Antarctic ice fish species (Chionodraco rastrospinosus, Cryodraco antarcticus, and Chaenodraco wilsoni). Stomach content analyses were conducted during bottom trawl surveys around the South Shetland Islands in 1998, 2001, 2002, 2003, and 2007, the South Orkney Islands in 1999 and 2009, and off the north-western Antarctic Peninsula in 2002, 2006, and 2006/2007 in order to obtain further insight into the amount of food of Antarctic demersal fish consume during summer. Annual variation in food intake was comparatively low within an area in the krill-feeding species C. gunnari and C. wilsoni. Food intake was much more variable, by a factor of 2 or 3 among years and areas, in larger C. aceratus and C. antarcticus, which rely heavily on fish as their dietary source. Food consumption was intermediate in the two species P. georgianus and C. rastrospinosus, which rely on both krill and fish.     

Diet of two icefish species from the South Shetland Islands and Elephant Island,<Emphasis Type="Italic"> Champsocephalus gunnari</Emphasis> and<Emphasis Type="Italic"> Chaenocephalus aceratus</Emphasis>

The summer diet of two species of icefishes (Channichthyidae) from the South Shetland Islands and Elephant Island, Champsocephalus gunnari and Chaenocephalus aceratus, was investigated from 2001 to 2003. Champsocephalus gunnari fed almost exclusively on krill (Euphausia superba) in all years. The importance of other taxa (Themisto gaudichaudii, mysids, myctophids) in the diet was negligible. The average feeding rate of Champsocephalus gunnari inferred from an exponential gastric evacuation model was between 1.0 and 1.5% body weight per day. Most of the stomachs of Chaenocephalus aceratus were empty. Stomachs with food contained mainly krill, mysids and fish. Among the fish taken, locally abundant species formed the bulk of the diet: Gobionotothen gibberifrons in 2001, Lepidonotothen larseni and Champsocephalus gunnari in 2002 and L. larseni in 2003. An ontogenetic shift in feeding preference of Chaenocephalus aceratus was observed: fish smaller than 30 cm fed on krill and mysids, while larger animals relied primarily on fish.     

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.     

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     

Fish prey of Antarctic fur seals Arctocephalus gazella at Ile de Croy,Kerguelen

 The composition of Antarctic fur seal prey was assessed through analysis of scats collected in March 1994 on Ile de Croy, Iles Nuageuses. Fish remains predominated in samples, occurring in 95% of droppings. A total of 968 otoliths allowed the identification of 16 fish species. Myctophid fish (12 species) dominated the diet both by number (94% of the otoliths) and by fish reconstituted mass (76%). Three fish species constituted together 87% of the reconstituted mass: the myctophids Gymnoscopelus nicholsi (52%) and G. piabilis (12%), and the channichthyid Champsocephalus gunnari (23%). Prey distribution suggests that during late summer seals forage in upper slope waters in the northeast of the Kerguelen Archipelago. Received:1 March 1996/Accepted: 20 May 1996     

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     

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 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.     

Sexual differences in the diet of king cormorants at Macquarie Island

Sexual differences in the diet of king cormorants,Phalacrocorax albiventer, were investigated during the 1993/1994 austral summer at Macquarie Island. The major food items, identified by otoliths in regurgitations, were demersal fish; fish mass consumed could be estimated using a wet mass-otolith length relationship. Two fish species,Paranotothenia magellanica andHarpagifer georgianus, constituted 98% of the wet mass (male and female cormorants combined). Estimated individual fish mass ofParanotothenia magellanica (19.6±11.6 g) was greater than that ofH. georgianus (2.8±1.3 g). Total wet mass of food and number of fish in regurgitations did not differ statistically between the sexes of cormorants. However, males tended to feed on larger fish than did females.     

Identifying patterns in the diet of mackerel icefish (<Emphasis Type="Italic">Champsocephalus gunnari)</Emphasis> at South Georgia using bootstrapped confidence intervals of a dietary index

Ontogenetic, inter-annual and regional variations in diet were investigated for mackerel icefish, Champsocephalus gunnari, in three successive summer seasons around South Georgia. Stomach contents from 2239 C. gunnari (130–560 mm total length) were examined. A bootstrapping technique was used to calculate confidence intervals for an index of relative importance of prey categories (% IRI_DC). Diet varied significantly between years and age classes but there was little regional difference in diet. In general, diet was dominated by krill, Euphausia superba and by the amphipod Themisto gaudichaudii. Smaller (younger) fish tended to prey on a higher proportion of T. gaudichaudii and small euphausiids such as Thysanoessa sp. and took smaller quantities of E. superba. In a season of poor krill availability (summer of 2003–2004) the proportion of krill in the diet, stomach fullness and fish condition (indicated by length–weight relationships) were significantly lower than in the other summer seasons. A large reduction (>80%) in the estimated annual (2005) biomass of the C. gunnari stock directly followed the season of poor krill availability. This decline was largely because of mortality of 2+ and 3+ fish, which were more krill dependent than 1+ fish. Younger fish appear to have survived, leading to an increase in the estimated population biomass in 2006.     

Analysis of stomach contents from the black-browed albatross, <Emphasis Type="Italic">Thalassarche melanophris</Emphasis>, on the Coast of Rio Grande do Sul,Southern Brazil

The black-browed albatross, Thalassarche melanophris, travels along the southern coast of Brazil during migration. Their numbers appear to have decreased in recent years. From July 1997 to 1998, we carried out monthly surveys on the coast of Rio Grande do Sul, Southern Brazil, to count albatrosses in the area and identify ingested items. We found 85 dead specimens of T. melanophris, from which 35 stomachs were collected and the contents analyzed. Nearly 6% of the stomachs contained fish, 32% contained cephalopods and 9% contained crustaceans. About 69% of the stomachs contained nematodes and nematomorphs. The highest abundance values found per stomach was 182 nematode individuals and 223 nematomorph individuals. Synthetic materials were present in 29% of the stomachs analyzed, comprising thermoplastic, nylon, rubber and metal wire.     

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.     

Muscle fibre types and metabolism in post-larval and adult stages of notothenioid fish

Summary A histochemical study was carried out on muscle fibre types in the myotomes of post-larval and adult stages of seven species of notothenioid fish. There was little interspecific variation in the distribution of muscle fibre types in post-larvae. Slow fibres (diameter range 15–60 m) which stained darkly for succinic dehydrogenase activity (SDHase) formed a superficial layer 1–2 fibres thick around the entire lateral surface of the trunk. In all species a narrow band of very small diameter fibres (diameter range 5–62 m), with only weak staining activity, occurred between the skin and slow fibre layer. These have the characteristics of tonic fibres found in other teleosts. The remainder of the myotome was composed of fast muscle fibres (diameter range 9–75 m), which stain weakly for SDHase, -glycerophosphate dehydrogenase, glycogen and lipid. Slow muscle fibres were only a minor component of the trunk muscles of adult stages of the pelagic species Champsocephalus gunnari and Pseudochaenichthys georgianus, consistent with a reliance on pectoral fin swimming during sustained activity. Of the other species examined only Psilodraco breviceps and Notothenia gibberifrons had more than a few percent of slow muscle in the trunk (20%–30% in posterior myotomes), suggesting a greater involvement of sub-carangiform swimming at cruising speeds. The ultrastructure of slow fibres from the pectoral fin adductor and myotomal muscles of a haemoglobinless (P. georgianus) and red-blooded species (P. breviceps), both active swimmers, were compared. Fibres contained loosely packed, and regularly shaped myofibrils numerous mitochondria, glycogen granules and occasional lipid droplets. Mitochondria occupied >50% of fibre volume in the haemoglobinless species P. georgianus, each myofibril was surrounded by one or more mitochondria with densely packed cristae. No significant differences, however, were found in mean diameter between fibres from red-blooded and haemoglobinless species. The activities of key enzymes of energy metabolism were determined in the slow (pectoral) and fast (myotomal) muscles of N. gibberifrons. In contrast to other demersal Antarctic fish examined, much higher glycolytic activities were found in fast muscle fibres, probably reflecting greater endurance during burst swimming.     

The diet of Antarctic fur sealsArctocephalus gazella during the breeding season at South Georgia

 The diet of lactating female Antarctic fur seals Arctocephalus gazella at South Georgia was investigated during the pup-rearing period (January/March) of 1991–1994. Antarctic krill Euphausia superba was the main prey item, occurring in 88% of all scats (n=497), whereas fish occurred in 47% and squid in 5% of all scats. There was considerable intra- and inter-annual variation in the characteristics of krill taken by fur seals. The distribution of krill sizes taken suggests that fur seals are not actively selecting particular sizes of krill and, therefore, that the krill in the diet reflects the krill available around South Georgia. The absence of group 3 krill (44–48 mm in length) in the South Georgia area, as indicated by their absence in the diet of seals, is suggested as a possible reason for low availability of krill and the subsequent reproductive failure among krill predators. The frequency of occurrence of fish was much higher than in previous studies; the pattern of fish consumption showed a consistent seasonal pattern in 3 of the 4 years studied. Of the total number of the myctophid Protomyctophum choriodon, the most numerous fish taxon, 98% were taken between early February and the middle of March. Champsocephalus gunnari and Lepidonotothen larseni agg., which both feed on krill, dominated the fish component of the diet outside this period and together constituted 94% of the total estimated biomass of fish consumed. The intra- and inter-annual variability in the diet of Antarctic fur seals emphasise the need for diet studies to be conducted during the entire pup-rearing periods of several years. Received: 15 March 1995/Accepted: 13 May 1995     

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.     

A comparative study of diet in three sympatric populations of<Emphasis Type="Italic"> Patagonotothen</Emphasis> species (Pisces: Nototheniidae)

The diet of three Patagonotothen species was investigated from the bycatch of near-bottom trawls for squid on the southern Falkland Island shelf. Patagonotothen ramsayi had a moderately diverse diet that was mostly zooplankton, especially amphipods, but included benthos and discards from the fishery, especially squid. P. guntheri also had a moderately diverse zooplankton diet, especially copepods, and benthos and squid damaged by the fishery. The diet of P. tessellata was the most diverse and was mainly benthos, including damaged scallops, with some euphausiids and amphipods. Feeding niche overlap of fish between 10 and 20 cm TL for the three species was between 5.8 and 9.2% for each pair of species, indicating relatively little overlap even though all species scavenged on organisms made available by the fishery. Thus the diet of these three Patagonotothen species indicates substantial feeding niche separation, and that the three species are moderately specialised feeders.