Energy costs of incubation in the Wandering Albatross Diomedea exulans

The role of the sexes in incubation, their changes in body weight and associated energy costs were studied in the Wandering Albatross at South Georgia. Males incubate significantly more than females (54%, range 36–68%) and lose weight during incubation shifts at a greater absolute, but smaller proportional, rate. On average this results in both sexes losing equal proportions of body weight during typical shifts. Energy costs of incubation are calculated to lie between 1.2 and 2.0 times basal metabolism, and probably much closer to the former.
All males and most females gained weight while at sea between shifts, most males doing so faster than females. On average both sexes are able at least to maintain body weight during the incubation period. More extreme individuals, however, may end incubation in poor condition and can probably redress this only by neglecting their chick. Chicks that died were more likely to have had parents with a less equal division of incubation duties.     

Factors determining nest-site selection of surface-nesting seabirds: a case study on the world's largest pelagic bird,the Wandering Albatross (Diomedea exulans)

Several factors may drive bird nest-site selection, including predation risk, resource availability, weather conditions and interaction with other individuals. Understanding the drivers affecting where birds nest is important for conservation planning, especially where environmental change may alter the distribution of suitable nest-sites. This study investigates which environmental variables affect nest-site selection by the Wandering Albatross Diomedea exulans, the world's largest pelagic bird. Here, wind characteristics are quantitatively investigated as a driver of nest-site selection in surface-nesting birds, in addition to several topographical variables, vegetation and geological characteristics. Nest locations from three different breeding seasons on sub-Antarctic Marion Island were modelled to assess which environmental factors affect nest-site selection. Elevation was the most important determinant of nest-site selection, with Wandering Albatrosses only nesting at low elevations. Distance from the coast and terrain roughness were also important predictors, with nests more generally found close to the coast and in flatter terrain, followed by wind velocity, which showed a hump-shaped relationship with the probability of nest occurrence. Nests occurred more frequently on coastal vegetation types, and were absent from polar desert vegetation (generally above c. 500 m elevation). Of the variables that influence Wandering Albatross nest location, both vegetation type and wind characteristics are likely to be influenced by climate change, and have already changed over the last 50 years. As a result, the availability of suitable nest-sites needs to be considered in light of future climate change, in addition to the impacts that these changes will have on foraging patterns and prey distribution. More broadly, these results provide insights into how a wide range of environmental variables, including wind, can affect nest-site selection of surface-nesting seabirds.     

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.     

The diet of the Wandering Albatross Diomedea exulans at Subantarctic Marion Island

Summary The diet of the Wandering Albatross at Subantarctic Marion Island was studied by inducing recently fed chicks to regurgitate and by stomach flushing adults about to feed chicks. Liquid comprised 70.2% of stomach content mass recovered from chicks. Solid material comprised cephalopods (58.6% by mass), fish (36.5%) and crustacean, cetacean and seabird material as minor items. Twenty-three taxa of cephalopods were identified, the onychcteuthid squid Kondakovia longimana being the most important. Estimated average mass of squid was 694 g with a maximum of over 8 kg. Diet of the Wandering Albatross at Marion Island was broadly similar to that at other studied localities. The high proportion of cephalopods known to float after death in the diet, and the deep-water habits of the few fish identified, suggest that scavenging plays an important role in foraging behaviour.     

Individual quality and reproductive performance in the Grey-headed Albatross Diomedea chrysostoma

The reproductive performances of Grey-headed Albatrosses Diomedea chrysostoma with a previous record (≤5 years) of consistent success (≤70% chicks fledged from eggs laid—"top" birds) or failure (≤ 70% of attempts failed—"bottom" birds) were compared during 1993 -1995. In 1995, top birds arrived back at the colony significantly earlier, had significantly shorter first and second incubation shifts and hatched larger chicks which grew significantly faster than bottom birds. In 1994, top birds also had larger hatchlings with higher rates of growth than bottom birds. In 1994, top birds had significantly higher hatching, fledging and therefore overall breeding success than bottom birds; very similar trends were evident in 1993 and 1995. Chick-rearing success and all indices of chick growth suggested that food availability was high in 1995 (and 1993) and low in 1994. Therefore the superior performance of top birds was maintained in years of very different conditions, with the chick-rearing period particularly critical. A simple model (using published demographic parameters for Grey-headed Albatrosses on South Georgia) suggests that top birds would produce 2.5 more chicks over their lifetime than bottom birds. With the currently declining population numbers, the relative contribution of top birds to the next generation may be even greater than this.     

Growth and energy expenditure of Wandering Albatross Diomedea exulans chicks

Wandering Albatross Diomedea exulans chicks require 9–10 months to achieve adult body size at fledging, at which time they are also sexually size dimorphic. Because the developmental period spans the winter season, chicks must endure severe winter conditions and variability in provisioning effort by their parents. Thus chicks may adjust their rate of energy utilization to accommodate variations in provisioning, but this has not previously been studied. We followed longitudinally the changes in growth, body composition and oxygen consumption of 10 chicks from the end of the brooding period until fledging on the Crozet Islands. Body mass, culmen length and wing length were measured every 10 days and total body water (TBW) and resting metabolic rate (RMR) were measured monthly. Overall growth followed a logistic curve for all chicks, and sexual dimorphism in body mass appeared as early as the second month of measurements (males being heavier than females). Absolute TBW followed a logistic increase like that of body mass and was significantly higher in males owing to the difference in body mass. Conversely, mass-specific TBW (i.e. the proportion of body mass made up of water) did not differ significantly between male and female chicks. Absolute RMR peaked at 1.5 × adult basal metabolism in midwinter when chicks achieved maximum body mass, but decreased to adult levels by the time chicks fledged. The decrease in absolute RMR following attainment of peak mass is atypical of most seabird chicks (Procellariiformes) and may be explained partly by a reduction in size of the gut when parents reduce provisioning effort. The changes in mass-specific RMR did not differ between sexes but male chicks, being heavier, had higher absolute oxygen consumption and therefore greater energy requirements.     

Breeding biology and current status of the Amsterdam Island Albatross Diomedea amsterdamensis

A small population of the 'Great Albatrosses' has been discovered on Amsterdam Island (Indian Ocean) and described in 1983 as a new species: Diomedea amsterdamensis (Roux et al. 1983). We present here data on the biometry, plumage variability and preliminary results of a long-term study on the breeding cycle and population dynamics of this relict population. The current population is estimated at 65 birds including 21 breeding pairs.     

Demographic Responses to Oxidative Stress and Inflammation in the Wandering Albatross (Diomedea exulans)

One of the major challenges in ecological research is the elucidation of physiological mechanisms that underlie the demographic traits of wild animals. We have assessed whether a marker of plasma oxidative stress (TBARS) and plasma haptoglobin (protein of the acute inflammatory phase response) measured at time t predict five demographic parameters (survival rate, return rate to the breeding colony, breeding probability, hatching and fledging success) in sexually mature wandering albatrosses over the next four years (Diomedea exulans) using a five-year individual-based dataset. Non-breeder males, but not females, having higher TBARS at time t had reduced future breeding probabilities; haptoglobin was not related to breeding probability. Neither TBARS nor haptoglobin predicted future hatching or fledging success. Haptoglobin had a marginally positive effect on female survival rate, while TBARS had a marginally negative effect on return rate. Our findings do not support the role for oxidative stress as a constraint of future reproductive success in the albatross. However, our data point to a potential mechanism underlying some aspects of reproductive senescence and survival. Our results also highlight that the study of the consequences of oxidative stress should consider the life-cycle stage of an individual and its reproductive history.     

Diving behaviour of the Shy Albatross Diomedea cauta in Tasmania: initial findings and dive recorder assessment

The diving behaviour of the Shy Albatross Diomedea cauta was investigated using archival time-depth recorders (TDRs) and maximum depth gauges (MDGs). Data from birds carrying multiple devices and from diving simulations indicated that the degree of correspondence between TDRs and MDGs varied with the dive depth, duration and frequency, as well as with body placement. The MDGs were the most reliable when the diving depth was greater than 0.5 m, when the diving frequency was low and when gauges were placed on the birds' backs. The TDRs were used during late incubation and early chick rearing in 1994. Fifty-two dives (0.4 m) were recorded during 20 foraging trips of 15 individuals. The majority of dives were within the upper 3 m of the water column and lasted for less than 6 s. However, dives to 7.4 m and others lasting 19 s were recorded. The albatrosses dived between 07.00 h and 22.00 h, with peaks in their diving activity near midday and twilight. Mean diving depth varied throughout the day. with the deepest dives occurring between 10.00 h and 12.00 h. Two dive types were identified on the basis of the relationship between dive depth and descent rate. Plunge dives were short (5 s), and the birds reached a maximum depth of 2.9 m. Swimming dives were both longer and deeper. The characteristics of Shy Albatross plunge dives were similar to those of gannets Morus spp., which are known to be proficient plunge divers. Swimming dives suggest that Shy Albatrosses actively pursue prey underwater.     

Extra-pair paternity in the strongly monogamous Wandering Albatross Diomedea exulans has no apparent benefits for females

Although 92% of avian species are socially monogamous, extra-pair copulation (EPC), resulting in extra-pair paternity (EPP), is a common reproductive strategy in birds. Among seabirds, in which the rate of social monogamy reaches 100%, Procellariiformes (albatrosses and petrels) show low EPP rates, with the noticeable exception of the only albatross investigated in this regard, the Waved Albatross Phoebastria irrorata . This species, in which forced copulations are known to occur, showed a surprisingly high rate of EPP (25% of chicks). We investigate here EPP rates in another albatross species, the Wandering Albatross Diomedea exulans , subject to a demographic survey conducted for 38 years. We combined data on pair bonds with analysis of ten microsatellite loci and found that 10.7% of 75 chicks had an extra-pair sire. Although there was some evidence for inbreeding avoidance, within-pair and extra-pair chicks showed similar levels of heterozygosity, and the incidence of EPP was independent of age, experience or past reproductive success. Hence, we found no evidence that females benefit from EPCs. Owing to the male-biased sex ratio in adults, widowed and divorced males required more time to find a new mate (+28 and +72%, respectively) than did females. Combined with high sexual size dimorphism, this phenomenon might promote the forced copulations observed in this species. Our data therefore suggest that EPC is beneficial to unpaired males but occurs at random in females, consistent with the hypothesis that EPP results solely from forced EPCs. However, the importance of the latter for EPP and the part played by solitary males require further investigation.     

Reproductive endocrinology of the Wandering Albatross Diomedea exulans in relation to biennial breeding and deferred sexual maturity

The reproductive endocrinology of the Wandering Albatross Diomedea exulans was studied at South Georgia to investigate the potential endocrine correlates of biennial breeding and of the acquisition of sexual maturity. Gonads of breeding birds and of known-age immature birds of both sexes were examined by laparoscopy throughout the period that they were at the nest site. Blood samples, subsequently analysed to determine concentrations of luteinizing hormone (LH), prolactin, progesterone, testosterone and oestradiol-17/i, were obtained from samples of breeding birds of both sexes at regular intervals from first arrival until the chicks fledged nearly a year later. Before laying in December, breeding birds had mature testes and ovarian follicles and high concentrations of LH, prolactin and sex steroids. Gonadal regression and a rapid drop in hormone levels (except for LH in females) occurred in early incubation (January). Testes (and follicles to a lesser extent) enlarged in mid-incubation, coinciding with high levels of LH and increases in prolactin and testosterone. Gonads finally regressed completely near hatching time. LH, prolactin and testosterone remained at low levels throughout chick rearing (April to November), but females had several periods of active progesterone and oestradiol secretion, and progesterone was detectable in males only late in the chick-rearing period. Although some changes in hormone levels are difficult to explain, the patterns are fairly typical of temperate birds. The persistence of progesterone secretion in both female breeders and non-breeding ‘immature’ birds is viewed as part of a mechanism inhibiting an ovary from becoming vitellogenic. Although testis size and testosterone concentrations increased with age in immature males (of ages 4–10 years), birds of 5 years and older are probably physiologically mature, even though breeding does not start until they are 7 years of age and only half an age group has bred by an age of 11 years. Immature females (of age 4–7 years) had undeveloped follicles, very low oestradiol concentrations but high progesterone levels, providing further support for the role of this hormone in inhibiting gonadotropin secretion. The condition of the female is therefore probably decisive in determining when a pair first attempts to breed but it is unknown what factors initiate normal ovarian development.     

Environmental modifications of a subantarctic peat-bog by the Wandering Albatross (Diomedea exulans): A preliminary study

Summary On Ile de la Possession, as on all the subantarctic islands, major compounds of organic nutrients are provided by avifauna. The presence of Wandering Albatrosses (Diomedea exulans) induces great perturbations in the environment at the site of their nest building activities: physical and chemical modifications of the soil (by removal of litter material for nest building, trampling and accumulated organic deposits) as well as microbiological, botanical and faunal changes. These perturbations are very localized in extent. The pedological changes have been described in terms of morphological modifications. The influence of Wandering Albatrosses on the vegetation and the invertebrate fauna has been assessed and shows a decrease in the number of plant species and an increase in the number of invertebrate species. The role of some of these as indicator species is discussed.     

Foraging movements of the Shy Albatross Diomedea cauta breeding in Australia; implications for interactions with longline fisheries

Satellite telemetry was used to identify the foraging zones of Shy Albatrosses Diomedea cauta breeding at two sites off Tasmania, Australia (Albatross Island in western Bass Strait and Pedra Branca to the south) to assess their level of interaction with longline fisheries. Adult birds from both colonies fed locally both in and outside the breeding season. Breeding birds from Albatross Island foraged over the Australian continental shelf or slope waters off northwest Tasmania, while those from Pedra Branca foraged between the colony and the southeastern edge of the continental shelf. The distances travelled by the birds and the duration of their foraging trips varied during the breeding cycle and tended to decrease as eggs approached hatching. Adults which were tracked near the end of the breeding season (March-April, n = 7 birds) deserted their chicks prematurely, and while dispersing further than incubating or brooding birds, they remained over the continental shelf and slope waters off southeast Australia. Home range analyses indicated 41% overlap between foraging zones of birds during successive breeding stages. Dispersal during the postbreeding period extended the foraging zones with less overlap between individuals (10% for Albatross Island and 19% for Pedra Branca). The recent contraction of the Japanese Southern Bluefin Tuna longline fishery to the south and east coasts of Tasmania has resulted in extensive overlap with adult Shy Albatrosses from Pedra Branca, but appears to pose a minimal threat to adult birds from Albatross Island. Coupled with the concomitant increase in the Australian domestic tuna longlining industry, adult Shy Albatrosses from southern Tasmania (Pedra Branca and the Mewstone) are vulnerable to incidental capture through out their annual cycle.     

Fly or die: the role of fat stores in the growth and development of Grey-headed Albatross Diomedea chrysostoma chicks

Chicks of albatrosses, like other Procellariiformes, become independent at a mass similar to their parents but during growth attain a peak mass some 30% or more greater, before losing mass prior to fledging. The current views are that this high peak mass represents chicks storing fat reserves as an energy sink, or as an insurance against periodic food scarcity, or as a Consequence of natural stochastic variation in provisioning rate. We analysed growth and body composition of Grey‐headed Albatross Diomedea chrysostoma chicks at Bird Island, South Georgia in 1984 and 1986, two years of very different food availability. In 1984 when overall breeding success was only 28% (the lowest in 20 years and less than halt that in 1986), chicks were significantly smaller in terms of peak mass (by 37%), primary length (by 25%), liver, lung, heart and kidney size (by 18–34%) and fat (by 75–80%) but not significantly different in terms of skeletal (tarsus, culmen, ulna, sternum) or muscle (pectoral, leg) size. Despite these differences, there were some important similarities in the patterns of growth in both years. Up to the attainment of peak mass, most of the growth of organs and of skeletal structures was completed and little fat was deposited. In the remaining part of the chick‐rearing period, feather growth and acquisition of fat stores were undertaken. Thus Grey‐headed Albatross chicks begin to acquire substantial fat stores only during the later part of the development period; this is contrary to the predictions of any of the existing hypotheses concerning provisioning patterns and the role of fat stores in Procellariiformes. We propose that the deposition of fat in the later stages of chick growth is an adaptation to: (a) ensure against energy demands and/or nutritional stress affecting the quality of flight feathers (many of which are not renewed for up to three years after fledging); and (b) provide an energy reserve for chicks to use in the critical period immediately after independence.     

Utilization of offal and discards from "finfish" trawlers around the Falkland Islands by the Black-browed Albatross Diomedea melanophris

Multinational fisheries operating in the vicinity of the Falkland Islands currently take c. 90,000 tonnes of true fish ("finfish" as opposed to squid) per annum, including hakes Merluccius spp., Southern Blue Whiting Micromesistius australis, Hoki Macruronus magel-lanicus and Red Cod Salilota australis, and generate substantial quantities of fisheries waste in the form of discards and offal. This paper examines the use made of this waste by scavenging Black-browed Albatrosses Diomedea melanophris which breed in the Falklands. The various types of waste produced are described and their consumption by scavenging albatrosses is quantified. The results indicate that Black-browed Albatrosses obtain c. 8000 tonnes of food per annum from this source, of which two-thirds is offal and the remainder whole discards. The energy content of this waste is equivalent to 4.4% of the estimated total annual energy requirements of the Falklands Black-browed Albatross population. However, as the fishery is a greater predator of finfish stocks than are the albatrosses, its long-term impact may be detrimental to these birds.     

Influence of adult breeding experience on growth and provisioning of Wandering Albatross Diomedea exulans chicks at South Georgia

For most seabirds, reproductive performance improves with age; in albatrosses this is thought not to be so (experience being acquired before starting breeding) but only one study (of chick growth in a single season at one site) has specifically addressed this. We compared the provisioning performance and growth rates of chicks of Wandering Albatrosses Diomedea exulans breeding for the first (IN), second and third (LE) and fourth or more times (EE) on Bird Island, South Georgia in the austral winters of 1996 and 1997. Eggs from EE adults were significantly heavier than the other two categories and these chicks had a greater mass and longer wings up to 160 days of age and longer culmen and tarsus up to 115 days old. However chicks from all categories fledged at the same average mass, size and age. No significant differences between categories in feeding frequency or meal size were detected but experienced adults made shorter long foraging trips and spent more time at the nest than less experienced birds. Adults that remained at the nest gave chicks smaller meals than those that left immediately after feeding the chick. Although provision of smaller but more frequent meals by experienced adults promotes more rapid chick growth, the resulting differences do not persist into the late chick-rearing period. Our results were very similar to those from Iles Crozet in the Indian Ocean, supporting the hypothesis that when Wandering Albatrosses start to breed they are fully competent foragers but that it takes a while, during early chick-rearing, for birds breeding for the first time to adapt to the additional demands of provisioning a chick.     

Attendance patterns and behaviour in relation to experience and pair-bond formation in the Wandering Albatross Diomedea exulans at South Georgia

Recruitment of Wandering Albatrosses Diomedea exulans to the breeding population at South Georgia took between 2 and 8 years after they first returned to their natal colony. In successive seasons, from first return to pairing, the date of arrival became earlier and the number of days spent ashore and the time spent interacting with other birds increased. Pairing birds arrived earlier and spent more time ashore than birds of similar experience which did not pair in that season. In the season following pairing they returned at the same time as breeding birds, but most did not breed; when ashore they spent much of their time alone or with their partner at the nest site. They left in mid-season before other non-breeders and bred the following season. Some birds accomplished this process by spending much time (50–60 days) ashore in two or three seasons but most birds spent a similar total time ashore spread over more seasons. Until the season prior to breeding, the number of birds of the opposite sex with whom interactions occurred was proportional to the amount of time spent ashore. There was, therefore, considerable scope for inter-individual assessment of potential partners before interactions were confined essentially to a single partner in the season before the first breeding attempt.     

Polychlorinated Biphenyls in the Plasma and Preen Oil of Black-Footed Albatross (Diomedea nigripes) Chicks and Adults on Midway Atoll,North Pacific Ocean

Polychlorinated biphenyls (PCBs) are ubiquitous in the environment. Midway Atoll, located in the North Pacific Ocean, was occupied by the military during and after World War II. However, Midway Atoll has become a national wildlife refuge and home to many different seabirds today, including the black-footed albatross (Diomedea nigripes) (BFAL). The profiles and toxic equivalents (TEQ) of PCB congeners in the plasma and preen oil of BFAL chicks and adults were determined in this study. The concentrations of the total PCBs in the plasma samples of chicks and adults collected in Midway Atoll ranged from 2.3 to 223.8 (mean 80.1) and 22.8 to 504.5 (mean 158.6) ng g^-1 (wet weight, ww), respectively. The TEQs ranged from 0.2 to 0.6 (mean 0.4) and 0.4 to 1.6 (mean 0.9) pg g^-1 ww, respectively, in the plasma samples of chicks and adults from Midway Atoll. The major congeners in the plasma samples of chicks and adults included PCBs 31, 87, 97, 99, 118, 138, 153, and 180, accounting for 70% of the total PCBs. The concentrations of the total PCBs in the adult preen oil samples ranged from 1693 to 39404 (mean 10122) ng g^-1 (ww), of which 97% were PCBs 105, 118, 128, 138, 153, 161, 172, and 183.