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.     

Quantitative analysis of the use of discards from squid trawlers by Black-browed Albatrosses Diomedea melanophris in the vicinity of the Falkland Islands

In the past decade, a major trawl fishery for the squid Loligo gahi has developed in the vicinity of Beauchêne Island, an internationally important breeding site for the Black-browed Albatross Diomedea melanophris. The breeding season diet of this albatross in the Falklands and its use of discards generated by the Loligo fishery were investigated. Albatross chicks are fed extensively on commercially exploited species of squid and fish including Loligo gahi and southern blue whiting Micromesistius australis. The quantity of waste generated by the Loligo fishery amounts to c. 5% of the reported catch and just over 50% of this waste, mainly Loligo and nototheniid fish, is scavenged by adult Black-browed Albatrosses. The total quantity scavenged during the chick rearing period amounts to 1000–2000 tonnes per year. This is equivalent to 10–15% of the total food requirement of the breeding Black-browed Albatross population on Beauchene Island during the period when the fishery is operating. Although the Loligo fishery currently provides a significant quantity of food to these albatrosses, its net effect may be detrimental to them, as it is a much greater predator of Loligo stocks than the albatrosses are estimated to have been prior to the fishery's development.     

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.     

Demographic parameters of black-browed albatrosses Thalassarche melanophris from the Falkland Islands

Black-browed albatrosses Thalassarche melanophris are currently classified as globally endangered. The most important populations of this species are believed to be declining due to, amongst other factors, unsustainable levels of incidental mortality in fishing gear. However, detailed demographic data are lacking for several critical populations, including the largest of all, nesting in the Falkland Islands. Here, we present data from the first Falkland Islands detailed demographic study (at New Island) and show that, from 2003 to 2009, the mean adult survival probability was 0.942 (95% CI: 0.930–0.952). Nesting frequency of adults is amongst the highest recorded for Thalassarche albatrosses and breeding success (0.564 chicks per egg) is within normal values. The nesting population in the intensively studied plots experienced an increase of 4% per year from 2004 to 2009. These results indicate that the Falklands population may not be as threatened as previously supposed, although studies from more sites and a longer time series are needed to confirm or refute this. The high survival rates may partly reflect recent efforts to mitigate bycatch made by the Falkland Islands and other fisheries in the region. The reinforcement of such initiatives may be critical to buffer the black-browed albatross population against ecosystem shifts and natural disasters (such as harmful algal blooms) that will likely become more frequent with ongoing global changes.     

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.     

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.     

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.     

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.     

Brood‐guarding duration in black‐browed albatrosses Thalassarche melanophris: temporal,geographical and individual variation

In birds, the period spent brooding or guarding young chicks is highly variable, but such variation has seldom been studied. Previous single‐year studies of Antarctic petrels Thalassoica antarctica and grey‐headed albatrosses Thalassarche chrysostoma revealed a pronounced seasonal decline in brood‐guarding duration and gave rise to the ‘synchronisation hypothesis’, which suggests that some of the variation in the length of the brood‐guarding stage is related to predictable seasonal changes in the risk of chick predation. We tested the predictions of this and three other hypotheses in a two‐site, four‐year study of the black‐browed albatross T. melanophris. The existence of a pronounced seasonal decline in brood‐guarding duration was apparent at both sites, and in years of contrasting food availability, providing further support for the ‘synchronisation hypothesis’. Alternative explanations for this pattern are that short brood‐guarding periods for late‐hatched chicks result from a seasonal decline in food availability or from the fact that early nesting birds are of higher individual quality. However, these explanations are at odds with the absence of a seasonal decline in early chick growth or in probability of chick survival. Furthermore, adult quality (measured as past reproductive performance) had a weak and inconsistent effect on the duration of brood‐guarding. Weather changes explained some of the variation in brood‐guarding, but there were no differences between regions of contrasting climates. Individual pairs displayed a degree of inter‐annual consistency in brood‐guarding duration and, at least in some years, longer brood‐guarding resulted in higher fledging probability. We speculate that a higher investment in brood‐guarding increases the cost of reproduction, which counteracts other selective pressures that would otherwise lead to longer brood‐guarding durations.     

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.     

The evolution of parent–offspring conflict over mate choice

In human societies, parents often have a strong influence on the mate choice of their offspring. Moreover, empirical studies show that conflict over mate choice between parents and offspring is widespread across human cultures. Here we provide the first theoretical investigation into this conflict, showing that it may result from an underlying evolutionary conflict over parental resource distribution. We present a series of evolutionary simulations in which we gradually expand a standard model of sexual selection by the stepwise addition of elements of parental involvement. In our model, females obtain resources enhancing their fecundity from both their chosen mate and their parents. Potential mates differ in their ability to provide resources and may signal this ability. Both females and their parents can develop a preference for the signal, with both preferences influencing the realized mate choice of the female. Parents may differentially allocate resources among their daughters depending on the resource-provisioning abilities of their sons-in-law. When fecundity returns on investment are diminishing, we find that parents invest most in daughters whose mates provide few resources. Subsequently, the daughters evolve to exploit this allocation rule through their mate choice, which is not in the parents' best interests. This results in a conflict over mate choice between parents and their offspring, manifested as an on-going divergence of offspring and parental preferences. We predict that the conflict should be most pronounced when fathers, as opposed to mothers, control resource allocation.     

Step- and genetic parent/offspring relationships in a Caribbean village

Ethnographic data from a rural Trinidadian village were examined to test for differences in step-and genetic parent/offspring relationships. The data indicated that when both step- and genetic offspring are co-resident in the same household, fathers interact more frequently and less agonistically with genetic offspring than they do with step-offspring. Contrary to predictions from attachment theory, two possible mechanisms for paternal attachment, duration of co-residence and co-residence at birth, are associated with lower rates of interaction, and higher rates of agonistic interaction with stepoffspring. The data also indicate gender differences in step- and genetic parent/ offspring relationships, higher rates of “fosterage” for stepoffspring, higher rates of emigration from the village for stepoffspring, and lower reproductive success for individuals raised by a stepparent.     

Lower birth weight of offspring born after self-poisoning of parent.

Birth weight was evaluated in 777 and 217 livebirths, respectively, of index females and female partners of index males born before and after severe self-poisoning with drugs. Birth weight was also evaluated in matched controls. Babies born to index females months or years after an attempted self-poisoning were found to have a lower birth weight than before this suicide attempt. The difference in the birth weight of subsequent pregnancies of index and control females was also highly significant. A similar trend was observed in livebirths of female partners of index males. However, the differences were not significant between previous and subsequent pregnancies and between index cases and matched controls.     

Assessing parent numbers from offspring genotypes: the importance of marker polymorphism

Methods to infer parent numbers from offspring genotypes either determine the minimum number of parents required to explain alleles and multilocus genotypes detected in the offspring or use models to incorporate information on population allele frequencies and allele segregation. Disparate results by different approaches suggest that one or perhaps all methods are subject to bias. Here, we investigate the performance of minimum parent number estimates, maximum likelihood, and Bayesian analyses (programs COLONY and PARENTAGE) with respect to marker information content in simulated data sets without knowledge of parental genotypes. Offspring families of different sizes were assumed to share one parent and to be sired by 1 or 5 additional parents. All methods committed large errors in terms of underestimation (minimum value) and overestimation (COLONY), or both (PARENTAGE) of parent numbers, unless the data were highly informative, and their relative performances depended on full-sib group sizes and sire numbers. Increasing the number of markers with low gene diversity (H(e) < or = 0.68) yielded only slow improvement of the results, but all 3 methods performed well with 5-7 markers of H(e) = 0.84. We emphasize the importance of high marker polymorphism for inferring parent numbers and individual parent contributions, as well as for the detection of monogamous reproduction.     

Transgenerational effects of parent and grandparent gender on offspring development in a biparental beetle species

Parental effects on offspring life-history traits are common and increasingly well-studied. However, the extent to which these effects persist into offspring in subsequent generations has received less attention. In this experiment, maternal and paternal effects on offspring and grand-offspring were investigated in the biparental burying beetle Nicrophorus vespilloides, using a split-family design. This allowed the separation of prenatal and postnatal transgenerational effects. Grandparent and parent gender were found to have a cumulative effect on offspring development and may provide a selection pressure on the division of parental investment in biparental species.