Foraging energetics of Grey-headed Albatrosses Diotnedea chrysostoma at Bird Island,South Georgia

At-sea metabolism (CO₂ production) and water turnover of six breeding Grey-headed Albatrosses Diomedea chrysostoma were measured, using the doubly labelled water method, at Bird Island, South Georgia, Mean food consumption (estimated from a water influx rate of 1.01 1 d^-1 and data on dietary composition) was 1200gd^-1 or 50.4 W. At-sea metabolism (derived from a rate of CO₂ production of 3.98 1 h^-1) was 27.7 W, 2.5 times the estimated basal metabolic rate (BMR). On average the birds ingested nearly twice as much food energy as they expended to obtain it. The metabolic rate during flight (estimated from at-sea metabolism and activity budget data) was 36.3 W (range 34.7–39.0 W) or 3.2 (range 3.0–3.4) times the predicted BMR. This is the lowest cost of flight yet measured, but consistent with the highly developed adaptations for economic flight shown by albatrosses. These results are briefly compared with data for other polar vertebrates (penguins, fur seals) exploiting similar prey.     

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.     

Foraging energetics of Grey-headed Albatrosses Diotnedea chrysostoma at Bird Island, South Georgia

At-sea metabolism (CO₂ production) and water turnover of six breeding Grey-headed Albatrosses Diomedea chrysostoma were measured, using the doubly labelled water method, at Bird Island, South Georgia, Mean food consumption (estimated from a water influx rate of 1.01 1 d^-1 and data on dietary composition) was 1200gd^-1 or 50.4 W. At-sea metabolism (derived from a rate of CO₂ production of 3.98 1 h^-1) was 27.7 W, 2.5 times the estimated basal metabolic rate (BMR). On average the birds ingested nearly twice as much food energy as they expended to obtain it. The metabolic rate during flight (estimated from at-sea metabolism and activity budget data) was 36.3 W (range 34.7–39.0 W) or 3.2 (range 3.0–3.4) times the predicted BMR. This is the lowest cost of flight yet measured, but consistent with the highly developed adaptations for economic flight shown by albatrosses. These results are briefly compared with data for other polar vertebrates (penguins, fur seals) exploiting similar prey.     

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.     

Population dynamics of Black-browed and Grey-headed Albatrosses Diomedea melanophrys and D. chrysostoma at Campbell Island, New Zealand, 1942–96

The numbers of Black-browed Albatrosses Diomedea melanophrys and Grey-headed Albatrosses D. chrysostoma at Campbell Island, New Zealand, have declined dramatically since the 1940s. Black-browed Albatross numbers went into a steep decline in the 1970s and, since at least 1984, have been increasing slightly at average rates of 1.1% and 2.1% per annum at two colonies. The long-term downward trend in numbers of the Grey-headed Albatross has continued into the 1990s, averaging annually between 3.0% and 4.8% per annum at different colonies. A demographic study carried out between 1984 and 1996 indicates that Black-browed and Grey-headed Albatrosses have similarly high annual adult survival rates (0.945 and 0.953, respectively). Black-browed Albatrosses breed for the first time at a younger average age than do Grey-headed Albatrosses (10 years and 13.5 years, respectively), have a higher average breeding success (0.663 compared with 0.397 for the latter species) and are annual breeders whereas Greyheaded Albatross show a typical biennial pattern of breeding. Both show low survival from fledging to first breeding; averaging 0.186 and 0.162 for Black-browed and Grey-headed Albatrosses, respectively. Both species are accidentally killed in the Japanese long-line fishery for tuna Thunnus sp. in the Australasian region. The steep decline of Black-browed Albatross numbers in the 1970s was concomitant with the development of this fishery in the foraging region of the Campbell Island birds. Currently, the slight increase in numbers is due to high adult survival rates and breeding success, and is coincident with a great reduction in long-line fishing. With stable and high adult survival rates, it is expected that future population trends will be mainly influenced by the recruitment rates. The continuous decline in Grey-headed Albatross numbers since the 1940s, before long-line fishing developed in this region, indicates that natural environmental processes contributed to the downward trend in breeding numbers. Modelling indicates that Grey-headed Albatross numbers will continue to decrease with the present demographic parameters. A comparison between the species breeding at different sites shows that differing environmental conditions influence demographic characteristics.     

Egg formation and the pre-laying period of Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma at Bird Island, South Georgia

Egg composition and factors influencing egg formation were studied in Black-browed and Grey-headed Albatrosses Diomedea melanophris and D. chrysostoma at Bird Island, South Georgia. At nests where eggs were laid, females arrived 6–7 days after males, stayed one day during which 96% of observed copulations occurred, then departed to sea for c. 16 days in D. chrysostoma, c. 10 days in D. melanophris , returning c. two days before laying. Yolk deposition, however, lasted 21 and 20 days, and started 32 and 29 days before laying, in D. chrysostoma and D. melanophris respectively. Therefore, it is probably initiated by environmental factors not by copulation. Egg, albumen and yolk mass are significantly greater in D. chrysostoma but the proportionate composition of the species' eggs is nearly identical. Reduced differences in chick mass at hatching may reflect the longer incubation period in D. chrysostoma or relate to subsequent differences in chick growth rate.     

Physiological conditions of parent and offspring Black-browed Albatrosses Thalassarche melanophris

Capsule: In Black-browed Albatrosses Thalassarche melanophris nutritional condition is correlated between parents and their offspring.

Aims: To test resource allocation hypotheses analysing the relationship between parental and offspring nutritional condition.

Methods: We measured blood chemistry parameters related with nutritional condition in 24 parents and their nestlings in a colony of Black-browed Albatrosses.

Results: There were no significant differences in blood parameters between sexes or location of the nest within the colony, neither among adults nor among nestlings. We found a significant positive correlation between parents and the nutritional condition of their offspring, measured as urea, uric acid and beta-hydroxybutyrate concentrations in blood.

Discussion: These relationships demonstrate that condition of the young seems to be merely a reflection of parental condition. An interesting relationship between alkaline phosphatase concentration in adults and nutritional condition of their nestlings was found, suggesting that age of the parents would be a key factor explaining quality of the nestling.     

Senescence effects in an extremely long-lived bird: the grey-headed albatross Thalassarche chrysostoma

Studies attempting to document reproductive or other pre-lethal senescence effects in wild birds typically face an array of problems, including flaws in statistical analyses, non-adaptive philopatry to deteriorating environments, confounding effects arising from cohort heterogeneity and differential death rates of phenotypes and the frequent pairing of old birds to younger mates. Furthermore, recent studies suggest that birds could maintain a high level of physical fitness until old age, before being struck by a catastrophic illness leading quickly to their demise. The presence of terminally ill individuals in most datasets (and their greater incidence in older age categories) may therefore provide a false impression of progressive senescence in cross-sectional analyses. This study was designed explicitly to avoid all the known pitfalls linked to the demonstration of progressive senescence in wild populations, and involved one of the very longest-lived bird species. We show that, during incubation, old (aged 35 years and over) male grey-headed albatrosses Thalassarche chrysostoma make longer foraging trips, and have lower daily mass gains, than experienced mid-aged individuals (aged up to 28 years). This is, to our knowledge, the first report documenting reduced foraging performance with old age. Hatching and breeding success of pairs composed of two old individuals were reduced in comparison to mid-aged pairs. Overall results were very similar when analyses were repeated using only individuals known to have survived 1 or 2 years beyond field measurements (hence probably not suffering from the effects of an advanced terminal illness). We conclude that extremely long-lived individuals usually experience some degree of general physical deterioration, leading to reduced foraging and breeding performance, long before their final demise.     

Reproductive endocrinology of the Black-browed albatross Diomedea melanophris and the Grey-headed albatross D. chrysostoma

Gonadal size and the circulating concentrations of two pituitary hormones (luteinizing hormone and prolactin) and three gonadal steroids (testosterone, progesterone and oestradiol-17β) were measured in two closely related Diomedea albatrosses at South Georgia. The Grey-headed albatross D. chrysostoma , if successful in rearing a chick, usually breeds biennially, whilst the Black-browed albatross D. melanophris normally breeds annually. Direct examination (by laparoscopy) of the gonads showed that the testes of both species underwent annual cycles, whilst endocrine data confirmed that those male Grey-headed albatrosses at the colony in the pre-laying period but not breeding in that year (having bred successfully the previous year) were apparently in full reproductive condition with elevated testosterone levels typical of breeding birds. However, the females of the two species differed markedly. Grey-headed albatrosses, in a year following successful breeding, had undeveloped ovaries with low levels of circulating oestradiol but high levels of progesterone, whereas the Black-browed albatrosses showed a pattern consistent with annual ovarian development. The profiles of gonadal steroids through the breeding season were similar for the males of both species but differences existed between the females. In the female Grey-headed albatrosses, transient peaks of progesterone were present throughout chick rearing but these were absent from Black-browed albatrosses. Prolactin had a similar profile in both species, with uniformly high levels throughout incubation and a rapid fall near the end of the brood-guard period. It is suggested that Grey-headed, like Black-browed, albatrosses are intrinsically annual breeders. However, if a female Grey-headed albatross breeds successfully in one year, then nutritional factors operate to ensure that in the following year the female does not show ovarian development, although the ovary is active in terms of progesterone secretion.     

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.     

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.     

Chick provisioning rates and growth in Blacklbrowed Albatross Diomedea melanophris and Grey-headed Albatross D. chrysostoma at Bird Island, South Georgia

We compared the parental division of labour and the pattern and rate of parental provisioning by two sympatric species of albatross of similar mass and breeding timetable but differing in diet and in the duration of chick‐rearing. Using electronic weighing platforms inside artificial nests, we recorded chick mass of Black‐browed Albatross and Grey‐headed Albatross at Bird Island, South Georgia every 10 minutes for both species in 1993 and 1994 and for each species in two other years between 1990 and 1996. The chick mass data (nearly one million weighings) were used to calculate meal mass (over 5000 meals) and intervals between meals. Adult birds were fitted with radio‐transmitters which allowed each meal to be allocated to the appropriate parent. The combination of meal mass and foraging trip duration were used to calculate provisioning rates for chicks and individual adults. Overall, Black‐browed Albatrosses delivered significantly lighter meals (569 g) than Grey‐headed Albatrosses (616 g) but more frequently (every 2.07 days and 2.50 days respectively). Thus combining foraging trip data for both parents, Black‐browed Albatross chicks received a meal every 1.22 days compared with 1.26 days for Greyheaded Albatross. These rates did not differ significantly. The contribution of each sex of each species in chick provisioning fluctuated between years, being similar in some years or biased towards males in others. Chicks of both species that failed to fledge received smaller, less frequent meals than successful chicks. In 1990 and 1994, Black‐browed Albatross chick provisioning rates were lower than in 1992 and 1993. In 1990, both meal mass and trip duration were affected, but only in 1994 was trip duration longer. Greyheaded Albatross chick provisioning rate was lower in 1994 than in other years but trip duration was longer. In each species, significant changes in meal mass and trip duration occurred within the chick‐rearing period. Chick provisioning rates invariably declined before chicks attained their peak mass. For both species, chick growth rates and peak and fledging mass, but not fledging age, were affected by differences in provisioning rate.     

Grey-headed Woodpecker,Picus canus,in north-eastern Turkey

Abstract

A new record of the Grey-headed Woodpecker in north-eastern Turkey shows, together with published results, that the species apparently is distributed in low density over the whole Black Sea region of Turkey.     

From the Eye of the Albatrosses: A Bird-Borne Camera Shows an Association between Albatrosses and a Killer Whale in the Southern Ocean

Albatrosses fly many hundreds of kilometers across the open ocean to find and feed upon their prey. Despite the growing number of studies concerning their foraging behaviour, relatively little is known about how albatrosses actually locate their prey. Here, we present our results from the first deployments of a combined animal-borne camera and depth data logger on free-ranging black-browed albatrosses (Thalassarche melanophrys). The still images recorded from these cameras showed that some albatrosses actively followed a killer whale (Orcinus orca), possibly to feed on food scraps left by this diving predator. The camera images together with the depth profiles showed that the birds dived only occasionally, but that they actively dived when other birds or the killer whale were present. This association with diving predators or other birds may partially explain how albatrosses find their prey more efficiently in the apparently ‘featureless’ ocean, with a minimal requirement for energetically costly diving or landing activities.     

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.     

An unusual source of apparent mitochondrial heteroplasmy: duplicate mitochondrial control regions in Thalassarche albatrosses

Molecular ecologists, in search of suitable molecular markers, frequently PCR-amplify regions of mitochondrial DNA from total DNA extracts. This approach, although common, is prone to the co-amplification of nuclear copies of transposed DNA sequences (numts), which can then generate apparent mitochondrial sequence heteroplasmy. In this study we describe the discovery of apparent mitochondrial sequence heteroplasmy in Thalassarche albatrosses but eliminate the possibility of true sequence heteroplasmy and numts and instead reveal the source of the apparent heteroplasmy to be a duplicated control region. The two control regions align easily but are not identical in sequence or in length. Comparisons of functionally significant conserved sequence blocks do not provide evidence of degeneration in either duplicate. Phylogenetic analyses of domain I of both control region copies in five Thalassarche species indicate that they are largely evolving in concert; however, a short section within them is clearly evolving independently. To our knowledge this is the first time contrasting evolutionary patterns have been reported for duplicate control regions. Available evidence suggests that this duplication may be taxonomically widespread, so the results presented here should be considered in future evolutionary studies targeting the control region of all Procellariiformes and potentially other closely related avian groups.     

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.