Mercury Concentrations in Feathers of Albatrosses and Large Petrels at South Georgia: Contemporary Patterns and Comparisons with Past Decades.

Mercury (Hg) is an environmental contaminant that can negatively impact the health of humans and wildlife. Albatrosses and large petrels show some of the highest levels of Hg contamination among birds, with potential repercussions for reproduction and survival. Here, body feather total Hg (THg) concentrations were determined in breeding adults of five species of albatrosses and large petrels in the foraging guild at South Georgia during the mid-2010s. We tested the effects of species, sex and trophic ecology (inferred from stable isotopes) on THg concentrations and compared our results with published values from past decades. Feather THg concentrations differed significantly among species (range: 1.9-49.6 µg g-1 dw), and were highest in wandering albatrosses Diomedea exulans, intermediate in black-browed albatrosses Thalassarche melanophris and northern giant petrels Macronectes halli, and lowest in southern giant petrels M. giganteus and white-chinned petrels Procellaria aequinoctialis. Females were more contaminated than males in all species, potentially due to differences in distributions and diet composition. Across species, THg concentrations were not correlated with feather δ13C or δ15N values, implying that species effects (e.g., breeding and moulting frequencies) may be more important than trophic effects in explaining feather THg concentrations in this foraging guild. Within species, the only significant correlation was between THg and δ13C in wandering albatrosses, which could reflect higher Hg exposure in subtropical waters. Comparisons with THg concentrations from past studies, which reflect contamination from 10 to > 60 years ago, revealed considerable annual variation and some evidence for increases over time for wandering and black-browed albatrosses since before 1950 and from the late 1980s, respectively.

Synchronous timing of return to breeding sites in a long-distance migratory seabird with ocean-scale variation in migration schedules.

BACKGROUND: Migratory birds generally have tightly scheduled annual cycles, in which delays can have carry-over effects on the timing of later events, ultimately impacting reproductive output. Whether temporal carry-over effects are more pronounced among migrations over larger distances, with tighter schedules, is a largely unexplored question. METHODS: We tracked individual Arctic Skuas Stercorarius parasiticus, a long-distance migratory seabird, from eight breeding populations between Greenland and Siberia using light-level geolocators. We tested whether migration schedules among breeding populations differ as a function of their use of seven widely divergent wintering areas across the Atlantic Ocean, Mediterranean Sea and Indian Ocean. RESULTS: Breeding at higher latitudes led not only to later reproduction and migration, but also faster spring migration and shorter time between return to the breeding area and clutch initiation. Wintering area was consistent within individuals among years; and more distant areas were associated with more time spent on migration and less time in the wintering areas. Skuas adjusted the period spent in the wintering area, regardless of migration distance, which buffered the variation in timing of autumn migration. Choice of wintering area had only minor effects on timing of return at the breeding area and timing of breeding and these effects were not consistent between breeding populations. CONCLUSION: The lack of a consistent effect of wintering area on timing of return between breeding areas indicates that individuals synchronize their arrival with others in their population despite extensive individual differences in migration strategies.

Deleterious effects of mercury contamination on immunocompetence, liver function and egg volume in an antarctic seabird.

Mercury (Hg) is a globally important pollutant that can negatively impact metabolic, endocrine and immune systems of marine biota. Seabirds are long-lived marine top predators and hence are at risk of bioaccumulating high Hg concentrations from their prey. Here, we measured blood total mercury (THg) concentrations and relationships with physiology and breeding parameters of breeding brown skuas (Stercorarius antarcticus) (n = 49 individuals) at Esperanza/Hope Bay, Antarctic Peninsula. Mean blood THg concentrations were similar in males and females despite the differences in body size and breeding roles, but differed between study years. Immune markers (hematocrit, Immunoglobulin Y [IgY] and albumin) were negatively correlated with blood THg concentrations, which likely indicates a disruptive effect of Hg on immunity. Alanine aminotransferase (GPT) activity, reflecting liver dysfunction, was positively associated with blood THg. Additionally, triacylglycerol and albumin differed between our study years, but did not correlate with Hg levels, and so were more likely to reflect changes in diet and nutritional status rather than Hg contamination. Egg volume correlated negatively with blood THg concentrations. Our study provides new insights into the sublethal effects of Hg contamination on immunity, liver function and breeding parameters in seabirds. In this Antarctic species, exposure to sublethal Hg concentrations reflects the short-term risks which could make individuals more susceptible to environmental stressors, including ongoing climatic changes.

Global assessment of marine plastic exposure risk for oceanic birds.

Plastic pollution is distributed patchily around the world's oceans. Likewise, marine organisms that are vulnerable to plastic ingestion or entanglement have uneven distributions. Understanding where wildlife encounters plastic is crucial for targeting research and mitigation. Oceanic seabirds, particularly petrels, frequently ingest plastic, are highly threatened, and cover vast distances during foraging and migration. However, the spatial overlap between petrels and plastics is poorly understood. Here we combine marine plastic density estimates with individual movement data for 7137 birds of 77 petrel species to estimate relative exposure risk. We identify high exposure risk areas in the Mediterranean and Black seas, and the northeast Pacific, northwest Pacific, South Atlantic and southwest Indian oceans. Plastic exposure risk varies greatly among species and populations, and between breeding and non-breeding seasons. Exposure risk is disproportionately high for Threatened species. Outside the Mediterranean and Black seas, exposure risk is highest in the high seas and Exclusive Economic Zones (EEZs) of the USA, Japan, and the UK. Birds generally had higher plastic exposure risk outside the EEZ of the country where they breed. We identify conservation and research priorities, and highlight that international collaboration is key to addressing the impacts of marine plastic on wide-ranging species.

Genomic insights into the secondary aquatic transition of penguins.

Penguins lost the ability to fly more than 60 million years ago, subsequently evolving a hyper-specialized marine body plan. Within the framework of a genome-scale, fossil-inclusive phylogeny, we identify key geological events that shaped penguin diversification and genomic signatures consistent with widespread refugia/recolonization during major climate oscillations. We further identify a suite of genes potentially underpinning adaptations related to thermoregulation, oxygenation, diving, vision, diet, immunity and body size, which might have facilitated their remarkable secondary transition to an aquatic ecology. Our analyses indicate that penguins and their sister group (Procellariiformes) have the lowest evolutionary rates yet detected in birds. Together, these findings help improve our understanding of how penguins have transitioned to the marine environment, successfully colonizing some of the most extreme environments on Earth.

Variation and correlation in the timing of breeding of North Atlantic seabirds across multiple scales.

Timing of breeding, an important driver of fitness in many populations, is widely studied in the context of global change, yet despite considerable efforts to identify environmental drivers of seabird nesting phenology, for most populations we lack evidence of strong drivers. Here we adopt an alternative approach, examining the degree to which different populations positively covary in their annual phenology to infer whether phenological responses to environmental drivers are likely to be (a) shared across species at a range of spatial scales, (b) shared across populations of a species or (c) idiosyncratic to populations. We combined 51 long-term datasets on breeding phenology spanning 50 years from nine seabird species across 29 North Atlantic sites and examined the extent to which different populations share early versus late breeding seasons depending on a hierarchy of spatial scales comprising breeding site, small-scale region, large-scale region and the whole North Atlantic. In about a third of cases, we found laying dates of populations of different species sharing the same breeding site or small-scale breeding region were positively correlated, which is consistent with the hypothesis that they share phenological responses to the same environmental conditions. In comparison, we found no evidence for positive phenological covariation among populations across species aggregated at larger spatial scales. In general, we found little evidence for positive phenological covariation between populations of a single species, and in many instances the inter-year variation specific to a population was substantial, consistent with each population responding idiosyncratically to local environmental conditions. Black-legged kittiwake Rissa tridactyla was the exception, with populations exhibiting positive covariation in laying dates that decayed with the distance between breeding sites, suggesting that populations may be responding to a similar driver. Our approach sheds light on the potential factors that may drive phenology in our study species, thus furthering our understanding of the scales at which different seabirds interact with interannual variation in their environment. We also identify additional systems and phenological questions to which our inferential approach could be applied.

Aquatic birds have middle ears adapted to amphibious lifestyles.

Birds exhibit wide variation in their use of aquatic environments, on a spectrum from entirely terrestrial, through amphibious, to highly aquatic. Although there are limited empirical data on hearing sensitivity of birds underwater, mounting evidence indicates that diving birds detect and respond to sound underwater, suggesting that some modifications of the ear may assist foraging or other behaviors below the surface. In air, the tympanic middle ear acts as an impedance matcher that increases sound pressure and decreases sound vibration velocity between the outside air and the inner ear. Underwater, the impedance-matching task is reversed and the ear is exposed to high hydrostatic pressures. Using micro- and nano-CT (computerized tomography) scans of bird ears in 127 species across 26 taxonomic orders, we measured a suite of morphological traits of importance to aerial and aquatic hearing to test predictions relating to impedance-matching in birds with distinct aquatic lifestyles, while accounting for allometry and phylogeny. Birds that engage in underwater pursuit and deep diving showed the greatest differences in ear structure relative to terrestrial species. In these heavily modified ears, the size of the input areas of both the tympanic membrane and the columella footplate of the middle ear were reduced. Underwater pursuit and diving birds also typically had a shorter extrastapedius, a reduced cranial air volume and connectivity and several modifications in line with reversals of low-to-high impedance-matching. The results confirm adaptations of the middle ear to aquatic lifestyles in multiple independent bird lineages, likely facilitating hearing underwater and baroprotection, while potentially constraining the sensitivity of aerial hearing.

Spatial and sex differences in mercury contamination of skuas in the Southern Ocean.

Antarctic marine ecosystems are often considered to be pristine environments, yet wildlife in the polar regions may still be exposed to high levels of environmental contaminants. Here, we measured total mercury (THg) concentrations in blood samples from adult brown skuas Stercorarius antarcticus lonnbergi (n = 82) from three breeding colonies south of the Antarctic Polar Front in the Southern Ocean (southwest Atlantic region): (i) Bahía Esperanza/Hope Bay, Antarctic Peninsula; (ii) Signy Island, South Orkney Islands; and, (iii) Bird Island, South Georgia. Blood THg concentrations increased from the Antarctic Peninsula towards the Antarctic Polar Front, such that Hg contamination was lowest at Bahía Esperanza/Hope Bay (mean ± SD, 0.95 ± 0.45 µg g-1 dw), intermediate at Signy Island (3.42 ± 2.29 µg g-1 dw) and highest at Bird Island (4.47 ± 1.10 µg g-1 dw). Blood THg concentrations also showed a weak positive correlation with δ15N values, likely reflecting the biomagnification process. Males had higher Hg burdens than females, which may reflect deposition of Hg into eggs by females or potentially differences in their trophic ecology. These data provide important insights into intraspecific variation in contamination and the geographic transfer of Hg to seabirds in the Southern Ocean.

Independent evolution of intermediate bill widths in a seabird clade.

Interspecific introgression can occur between species that evolve rapidly within an adaptive radiation. Pachyptila petrels differ in bill size and are characterised by incomplete reproductive isolation, leading to interspecific gene flow. Salvin's prion (Pachyptila salvini), whose bill width is intermediate between broad-billed (P. vittata) and Antarctic (P. desolata) prions, evolved through homoploid hybrid speciation. MacGillivray's prion (P. macgillivrayi), known from a single population on St Paul (Indian Ocean), has a bill width intermediate between salvini and vittata and could also be the product of interspecies introgression or hybrid speciation. Recently, another prion population phenotypically similar to macgillivrayi was discovered on Gough (Atlantic Ocean), where it breeds 3 months later than vittata. The similarity in bill width between the medium-billed birds on Gough and macgillivrayi suggest that they could be closely related. In this study, we used genetic and morphological data to infer the phylogenetic position and evolutionary history of P. macgillivrayi and the Gough medium-billed prion relative other Pachyptila taxa, to determine whether species with medium bill widths evolved through common ancestry or convergence. We found that Gough medium-billed prions belong to the same evolutionary lineage as macgillivrayi, representing a new population of MacGillivray's prion that originated through a colonisation event from St Paul. We show that macgillivrayi's medium bill width evolved through divergence (genetic drift) and independently from that of salvini, which evolved through hybridisation (gene flow). This represents the independent convergence towards a similarly medium-billed phenotype. The newly discovered MacGillivray's prion population on Gough is of utmost conservation relevance, as the relict macgillivrayi population in the Indian Ocean is very small.

Genetic diversity and structure of captive gentoo penguin populations in Japan.

Until the last decade, gentoo penguins were usually split into two subspecies, northern gentoo penguins (Pygoscelis papua papua) breeding in the Falkland Islands, South Georgia, and other subantarctic islands and southern gentoo penguins (P. papua ellsworthi) breeding in the South Sandwich, South Orkney and South Shetland islands, and Antarctic Peninsula. Recent genetics research, however, suggests that the population at South Georgia is much more closely related to those further south and should be included in P. papua ellsworthi. In Japanese zoos and aquariums, captive breeding of gentoo penguins is conducted separately in three populations: "Captive-South Georgia," originating from South Georgia, "Captive-South Shetlands," originating from South Shetlands, and "Captive-Unknown," originating from at least one founder of unknown subspecies. The aims of the present study were to investigate the genetic diversity and differentiation of these captive populations using microsatellite analysis. Genetic diversity in each captive population was similar to that found in the wild, although they had much lower contemporary effective population sizes. Pairwise genetic differentiation indexes (FST ) among the three captive populations were as follows: 0.0309 ("Captive-South Georgia" and "Captive-Unknown"), 0.1094 ("Captive-South Georgia" and "Captive-South Shetlands"), and 0.1214 ("Captive-South Shetlands" and "Captive-Unknown"). Using Bayesian clustering, there was relatively high genetic differentiation between the "Captive-South Shetlands" group, which formed a distinct cluster, and individuals of the "Captive-Unknown" group, which were assigned to clusters in common with "Captive-South Georgia." The results from the present study are useful for future management of captive gentoo penguin populations in Japan.

Untangling local and remote influences in two major petrel habitats in the oligotrophic Southern Ocean.

Ocean circulation connects geographically distinct ecosystems across a wide range of spatial and temporal scales via exchanges of physical and biogeochemical properties. Remote oceanographic processes can be especially important for ecosystems in the Southern Ocean, where the Antarctic Circumpolar Current transports properties across ocean basins through both advection and mixing. Recent tracking studies have indicated the existence of two large-scale, open ocean habitats in the Southern Ocean used by grey petrels (Procellaria cinerea) from two populations (i.e., Kerguelen and Antipodes islands) during their nonbreeding season for extended periods during austral summer (i.e., October to February). In this work, we use a novel combination of large-scale oceanographic observations, surface drifter data, satellite-derived primary productivity, numerical adjoint sensitivity experiments, and output from a biogeochemical state estimate to examine local and remote influences on these grey petrel habitats. Our aim is to understand the oceanographic features that control these isolated foraging areas and to evaluate their ecological value as oligotrophic open ocean habitats. We estimate the minimum local primary productivity required to support these populations to be much <1% of the estimated local primary productivity. The region in the southeast Indian Ocean used by the birds from Kerguelen is connected by circulation to the productive Kerguelen shelf. In contrast, the region in the south-central Pacific Ocean used by seabirds from the Antipodes is relatively isolated suggesting it is more influenced by local factors or the cumulative effects of many seasonal cycles. This work exemplifies the potential use of predator distributions and oceanographic data to highlight areas of the open ocean that may be more dynamic and productive than previously thought. Our results highlight the need to consider advective connections between ecosystems in the Southern Ocean and to re-evaluate the ecological relevance of oligotrophic Southern Ocean regions from a conservation perspective.

Plumage development and environmental factors influence surface temperature gradients and heat loss in wandering albatross chicks.

Young birds in cold environments face a range of age-specific thermal challenges. Studying the thermal biology of young birds throughout ontogeny may further our understanding of how such challenges are met. We investigated how age and environmental parameters influenced surface temperature gradients across various body regions of wandering albatross (Diomedea exulans) chicks on Bird Island, South Georgia. This study was carried out over a 200 d period during the austral winter, from the end of the brood-guard period until fledging, bridging a gap in knowledge of surface temperature variation and heat loss in developing birds with a long nestling stage in severe climatic conditions. We found that variation in surface temperature gradients (i.e. the difference between surface and environmental temperature) was strongly influenced by chick age effects for insulated body regions (trunk), with an increase in the surface temperature gradient that followed the progression of plumage development, from the second set of down (mesoptiles), to final chick feathers (teleoptiles). Environmental conditions (primarily wind speed and relative humidity) had a stronger influence on the gradients in uninsulated areas (eye, bill) than insulated regions, which we interpret as a reflection of the relative degree of homeothermy exhibited by chicks of a given age. Based on biophysical modelling, total heat loss of chicks was estimated to increase linearly with age. However, mass specific heat loss decreased during the early stages of growth and then subsequently increased. This was attributed to age-related changes in feather growth and activity that increased surface temperature and, hence, metabolic heat loss. These results provide a foundation for further work on the effects of environmental stressors on developing chicks, which are key to understanding the physiological responses of animals to changes in climate in polar regions.

Global political responsibility for the conservation of albatrosses and large petrels.

Migratory marine species cross political borders and enter the high seas, where the lack of an effective global management framework for biodiversity leaves them vulnerable to threats. Here, we combine 10,108 tracks from 5775 individual birds at 87 sites with data on breeding population sizes to estimate the relative year-round importance of national jurisdictions and high seas areas for 39 species of albatrosses and large petrels. Populations from every country made extensive use of the high seas, indicating the stake each country has in the management of biodiversity in international waters. We quantified the links among national populations of these threatened seabirds and the regional fisheries management organizations (RFMOs) which regulate fishing in the high seas. This work makes explicit the relative responsibilities that each country and RFMO has for the management of shared biodiversity, providing invaluable information for the conservation and management of migratory species in the marine realm.

Hidden Markov models identify major movement modes in accelerometer and magnetometer data from four albatross species.

BACKGROUND: Inertial measurement units (IMUs) with high-resolution sensors such as accelerometers are now used extensively to study fine-scale behavior in a wide range of marine and terrestrial animals. Robust and practical methods are required for the computationally-demanding analysis of the resulting large datasets, particularly for automating classification routines that construct behavioral time series and time-activity budgets. Magnetometers are used increasingly to study behavior, but it is not clear how these sensors contribute to the accuracy of behavioral classification methods. Development of effective  classification methodology is key to understanding energetic and life-history implications of foraging and other behaviors. METHODS: We deployed accelerometers and magnetometers on four species of free-ranging albatrosses and evaluated the ability of unsupervised hidden Markov models (HMMs) to identify three major modalities in their behavior: 'flapping flight', 'soaring flight', and 'on-water'. The relative contribution of each sensor to classification accuracy was measured by comparing HMM-inferred states with expert classifications identified from stereotypic patterns observed in sensor data. RESULTS: HMMs provided a flexible and easily interpretable means of classifying behavior from sensor data. Model accuracy was high overall (92%), but varied across behavioral states (87.6, 93.1 and 91.7% for 'flapping flight', 'soaring flight' and 'on-water', respectively). Models built on accelerometer data alone were as accurate as those that also included magnetometer data; however, the latter were useful for investigating slow and periodic behaviors such as dynamic soaring at a fine scale. CONCLUSIONS: The use of IMUs in behavioral studies produces large data sets, necessitating the development of computationally-efficient methods to automate behavioral classification in order to synthesize and interpret underlying patterns. HMMs provide an accessible and robust framework for analyzing complex IMU datasets and comparing behavioral variation among taxa across habitats, time and space.

No evidence for fitness signatures consistent with increasing trophic mismatch over 30 years in a population of European shag Phalacrocorax aristotelis.

As temperatures rise, timing of reproduction is changing at different rates across trophic levels, potentially resulting in asynchrony between consumers and their resources. The match-mismatch hypothesis (MMH) suggests that trophic asynchrony will have negative impacts on average productivity of consumers. It is also thought to lead to selection on timing of breeding, as the most asynchronous individuals will show the greatest reductions in fitness. Using a 30-year individual-level dataset of breeding phenology and success from a population of European shags on the Isle of May, Scotland, we tested a series of predictions consistent with the hypothesis that fitness impacts of trophic asynchrony are increasing. These predictions quantified changes in average annual breeding success and strength of selection on timing of breeding, over time and in relation to rising sea surface temperature (SST) and diet composition. Annual average (population) breeding success was negatively correlated with average lay date yet showed no trend over time, or in relation to increasing SST or the proportion of principal prey in the diet, as would be expected if trophic mismatch was increasing. At the individual level, we found evidence for stabilising selection and directional selection for earlier breeding, although the earliest birds were not the most productive. However, selection for earlier laying did not strengthen over time, or in relation to SST or slope of the seasonal shift in diet from principal to secondary prey. We found that the optimum lay date advanced by almost 4 weeks during the study, and that the population mean lay date tracked this shift. Our results indicate that average performance correlates with absolute timing of breeding of the population, and there is selection for earlier laying at the individual level. However, we found no fitness signatures of a change in the impact of climate-induced trophic mismatch, and evidence that shags are tracking long-term shifts in optimum timing. This suggests that if asynchrony is present in this system, breeding success is not impacted. Our approach highlights the advantages of examining variation at both population and individual levels when assessing evidence for fitness impacts of trophic asynchrony.

Mercury exposure in an endangered seabird: long-term changes and relationships with trophic ecology and breeding success.

Mercury (Hg) is an environmental contaminant which, at high concentrations, can negatively influence avian physiology and demography. Albatrosses (Diomedeidae) have higher Hg burdens than all other avian families. Here, we measure total Hg (THg) concentrations of body feathers from adult grey-headed albatrosses (Thalassarche chrysostoma) at South Georgia. Specifically, we (i) analyse temporal trends at South Georgia (1989-2013) and make comparisons with other breeding populations; (ii) identify factors driving variation in THg concentrations and (iii) examine relationships with breeding success. Mean ± s.d. feather THg concentrations were 13.0 ± 8.0 µg g-1 dw, which represents a threefold increase over the past 25 years at South Georgia and is the highest recorded in the Thalassarche genus. Foraging habitat, inferred from stable isotope ratios of carbon (δ13C), significantly influenced THg concentrations-feathers moulted in Antarctic waters had far lower THg concentrations than those moulted in subantarctic or subtropical waters. THg concentrations also increased with trophic level (δ15N), reflecting the biomagnification process. There was limited support for the influence of sex, age and previous breeding outcome on feather THg concentrations. However, in males, Hg exposure was correlated with breeding outcome-failed birds had significantly higher feather THg concentrations than successful birds. These results provide key insights into the drivers and consequences of Hg exposure in this globally important albatross population.

Long-term changes in habitat and trophic level of Southern Ocean squid in relation to environmental conditions.

Long-term studies of pelagic nekton in the Southern Ocean and their responses to ongoing environmental change are rare. Using stable isotope ratios measured in squid beaks recovered from diet samples of wandering albatrosses Diomedea exulans, we assessed decadal variation (from 1976 to 2016) in the habitat (δ13C) and trophic level (δ15N) of five important Southern Ocean squid species in relation to indices of environmental conditions-Southern Oscillation Index (SOI) and Southern Annular Mode (SAM). Based on δ13C values, corrected for the Suess effect, habitat had changed over the last 50 years for Taonius sp. B (Voss), Gonatus antarcticus, Galiteuthis glacialis and Histioteuthis atlantica but not Moroteuthopsis longimana. By comparison, mean δ15N values were similar across decades for all five species, suggesting minimal changes in trophic levels. Both SAM and SOI have increased in strength and frequency over the study period but, of the five species, only in Taonius sp. B (Voss) did these indices correlate with, δ13C and δ15N values, indicating direct relationships between environmental conditions, habitat and trophic level. The five cephalopod species therefore changed their habitats with changing environmental conditions over the last 50 years but maintained similar trophic levels. Hence, cephalopods are likely to remain important prey for top predators in Southern Ocean food webs, despite ongoing climate change.

Sex-specific effects of wind on the flight decisions of a sexually dimorphic soaring bird.

In a highly dynamic airspace, flying animals are predicted to adjust foraging behaviour to variable wind conditions to minimize movement costs. Sexual size dimorphism is widespread in wild animal populations, and for large soaring birds which rely on favourable winds for energy-efficient flight, differences in morphology, wing loading and associated flight capabilities may lead males and females to respond differently to wind. However, the interaction between wind and sex has not been comprehensively tested. We investigated, in a large sexually dimorphic seabird which predominantly uses dynamic soaring flight, whether flight decisions are modulated to variation in winds over extended foraging trips, and whether males and females differ. Using GPS loggers we tracked 385 incubation foraging trips of wandering albatrosses Diomedea exulans, for which males are c. 20% larger than females, from two major populations (Crozet and South Georgia). Hidden Markov models were used to characterize behavioural states-directed flight, area-restricted search (ARS) and resting-and model the probability of transitioning between states in response to wind speed and relative direction, and sex. Wind speed and relative direction were important predictors of state transitioning. Birds were much more likely to take off (i.e. switch from rest to flight) in stronger headwinds, and as wind speeds increased, to be in directed flight rather than ARS. Males from Crozet but not South Georgia experienced stronger winds than females, and males from both populations were more likely to take-off in windier conditions. Albatrosses appear to deploy an energy-saving strategy by modulating taking-off, their most energetically expensive behaviour, to favourable wind conditions. The behaviour of males, which have higher wing loading requiring faster speeds for gliding flight, was influenced to a greater degree by wind than females. As such, our results indicate that variation in flight performance drives sex differences in time-activity budgets and may lead the sexes to exploit regions with different wind regimes.

Introduced species and extreme weather as key drivers of reproductive output in three sympatric albatrosses.

Invasive species present a major conservation threat globally and nowhere are their affects more pronounced than in island ecosystems. Determining how native island populations respond demographically to invasive species can provide information to mitigate the negative effects of invasive species. Using 20 years of mark-recapture data from three sympatric species of albatrosses (black-browed Thalassarche melanophris, grey-headed T. chrysostoma, and light-mantled albatrosses Phoebetria palpebrata), we quantified the influence of invasive European rabbits Oryctolagus cuniculus and extreme weather patterns on breeding probability and success. Temporal variability in rabbit density explained 33-76% of the variability in breeding probability for all three species, with severe decreases in breeding probability observed after a lag period following highest rabbit numbers. For black-browed albatrosses, the combination of extreme rainfall and high rabbit density explained 33% of total trait variability and dramatically reduced breeding success. We showed that invasive rabbits and extreme weather events reduce reproductive output in albatrosses and that eliminating rabbits had a positive effect on albatross reproduction. This illustrates how active animal management at a local breeding site can result in positive population outcomes even for wide ranging animals like albatrosses where influencing vital rates during their at-sea migrations is more challenging.