A comparison of methods to evaluate energy expenditure of incubating wandering albatrosses

Measurements of incubation energetics can vary depending on the method used to measure metabolism of an incubating bird. Therefore, we evaluated the energy expenditure of six male and four female wandering albatrosses (Diomedea exulans Linnaeus) using doubly labeled water (DLW), the rate of mass loss, and estimates of metabolic water production derived from water influx rate (WIR). Incubation metabolic rates (IMR) determined with DLW ( 169 ± 21 kJ kg d SD) were significantly lower than estimates derived from mass loss ( 277 ± 46kJ kg d SD) and WIR ( males=289 ± 60 kJ kg d vs. females = 400 ± 69 kJ kg d SD). Estimates of IMR from f WIR were similar to IMR (305 ± 39 kJ kg d SD) determined by respirometry in a previous study, and IMR from DLW was similar to estimates based on heart rate (HR; 147 ± 26 kJ d SD) determined in another study. Ap- 147 26 plying the different measurements of IMR to construct an en-ergy budget, we estimate that a breeding pair of wandering albatrosses spends 124--234 MJ to incubate the egg for 78 d. Finally, IMRs determined with DLW and HR were similar

Corticosterone and foraging behaviour in a pelagic seabird

Because endocrine mechanisms are thought to mediate behavioral responses to changes in the environment, examining these mechanisms is essential for understanding how long-lived seabirds adjust their foraging decisions to contrasting environmental conditions in order to maximize their fitness. In this context, the hormone corticosterone (CORT) deserves specific attention because of its major connections with locomotor activities. We examined for the first time the relationships between individual CORT levels and measurements of foraging success and behavior using satellite tracking and blood sampling from wandering albatrosses (Diomedea exulans) before (pretrip CORT levels) and after (posttrip CORT levels) foraging trips during the incubation period. Plasma CORT levels decreased after a foraging trip, and the level of posttrip CORT was negatively correlated with individual foraging success, calculated as total mass gain over a foraging trip. Pretrip CORT levels were not linked to time spent at sea but were positively correlated with daily distance traveled and maximum range at sea. In this study, we were able to highlight the sensitivity of CORT levels to variation in energy intake, and we showed for the first time that individual CORT levels can be explained by variation in foraging success. Relationships between pretrip CORT levels and daily distance traveled and maximum range were independent of pretrip body mass, suggesting that slight elevations in pretrip CORT levels might facilitate locomotor activity. However, because both foraging behavior and pretrip CORT levels could be affected by individual quality, future experimental studies including manipulation of CORT levels are needed to test whether CORT can mediate foraging decisions according to foraging conditions

Wind field and sex constrain the flight speeds of central-place foraging albatrosses

By extracting energy from the highly dynamic wind and wave fields that typify pelagic habitats, albatrosses are able to proceed almost exclusively by gliding flight. Although energetic costs of gliding are low, enabling breeding albatrosses to forage hundreds to thousands of kilometers from their colonies, these and time costs vary with relative wind direction. This causes albatrosses in some areas to route provisioning trips to avoid headwind flight, potentially limiting habitat accessibility during the breeding season. In addition, because female albatrosses have lower wing loadings than males, it has been argued that they are better adapted to flight in light winds, leading to sexual segregation of foraging areas. We used satellite telemetry and immersion logger data to quantify the effects of relative wind speed, sex, breeding stage, and trip stage on the ground speeds (Vg) of four species of Southern Ocean albatrosses breeding at South Georgia. Vg was linearly related to the wind speed component in the direction of flight (Vwf), its effect being greatest on Wandering Albatrosses Diomedea exulans, followed by Black-browed Albatrosses Thalassarche melanophrys, Light-mantled Sooty Albatrosses Phoebatria palpebrata, and Gray-headed Albatrosses T. chrysostoma. Ground speeds at Vwf = 0 were similar to airspeeds predicted by aerodynamic theory and were higher in males than in females. However, we found no evidence that this led to sexual segregation, as males and females experienced comparable wind speeds during foraging trips. Black-browed, Gray-headed, and Light-mantled Sooty Albatrosses did not engage in direct, uninterrupted bouts of flight on moonless nights, but Wandering Albatrosses attained comparable Vg night and day, regardless of lunar phase. Relative flight direction was more important in determining Vg than absolute wind speed. When birds were less constrained in the middle stage of foraging trips, all species flew predominantly across the wind. However, in some instances, commuting birds encountered headwinds during outward trips and tail winds on their return, with the result that Vg was 1.0–3.4 m/s faster during return trips. This, we hypothesize, could result from constraints imposed by the location of prey resources relative to the colony at South Georgia or could represent an energy optimization strategy

Variability of resource partitioning in sympatric tropical boobies

International audienceInter- and intraspecific competition can lead to resource partitioning in sympatric species, processes likely affected by environmental productivity and population size. We investigated the foraging behaviour and diet of masked (Sula dactylatra) and red-footed (S. sula) boobies at Tromelin Island, western Indian Ocean, to examine the role of resource partitioning in the foraging strategies of these sympatric species in an extreme oligotrophic environment. We compared our results to published studies with differing environmental conditions or population sizes. We used GPS loggers and Argos transmitters to track foraging movements and used time-depth recorders to estimate dive depths. Masked boobies travelled further and at faster rates than red-footed boobies, and sexes did not differ in foraging behaviour. Based on randomization tests, the foraging range of each species (95% utilization distribution; UD) overlapped significantly. However, at core foraging areas (50% and 25% UD), interspecific segregation was greater than expected by chance alone. No intraspecific spatial segregation was detected between sexes. Environmental characteristics of area-restricted search zones differed between species, but not sexes; masked boobies utilized warmer, deeper, and less windy oceanic environments than red-footed boobies. Masked boobies attained greater diving depths than red-footed boobies and consumed primarily flying fish, whereas red-footed boobies consumed mostly squid. Red-footed and masked boobies breeding in the extreme oligotrophic environment near Tromelin demonstrated greater dietary partitioning differences in foraging ranges compared to a less oligotrophic environment. This suggests that environmental productivity may play a role in processes of resource partitioning in these sympatric species

Extreme variation in migration strategies between and within wandering albatross populations during their sabbatical year, and their fitness consequences

Migratory behavior, routes and zones used during the non-breeding season are assumed to have been selected to maximize fitness, and can lead to genetic differentiation. Yet, here we show that migration strategies differ markedly between and within two genetically similar populations of wandering albatross Diomedea exulans from the Crozet and Kerguelen archipelagos in the Indian Ocean. Wandering albatrosses usually breed biennially if successful, and during the sabbatical year, all birds from Kerguelen migrate to the Pacific Ocean, whereas most from Crozet are sedentary. Instead of taking the shortest routes, which would involve a return against headwinds, migratory birds fly with the westerly winds, requiring detours of 10,000 s km. In total, migrants circumnavigate Antarctica 2 to 3 times, covering more than 120,000 km in a single sabbatical year. Our results indicate strong links between migratory behavior and fitness; all birds from Kerguelen breed biennially, whereas a significant proportion of those from Crozet, especially females, are sedentary and breed in consecutive calendar years. To breed annually, these females temporarily change mate, but return to their original partner in the following year. This extreme variation in migratory behavior has important consequences in term of life history evolution and susceptibility to climate change and fisheries

Heart rate and energy expenditure of incubating wandering albatrosses: basal levels, natural variation, and the effects of human disturbance

We studied the changes in heart rate (HR) associated with metabolic rate of incubating and resting adult wandering albatrosses (Diomedea exulans) on the Crozet Islands. Metabolic rates of resting albatrosses fitted with external HR recorders were measured in a metabolic chamber to calibrate the relationship between HR and oxygen consumption (V̇O2) (V̇O2=0.074×HR+0.019, r2=0.567, P\u3c0.001, where V̇O2 is in ml kg–1 min–1 and HR is in beats min–1). Incubating albatrosses were then fitted with HR recorders to estimate energy expenditure of albatrosses within natural field conditions. We also examined the natural variation in HR and the effects of human disturbance on nesting birds by monitoring the changes in HR. Basal HR was positively related to the mass of the individual. The HR of incubating birds corresponded to a metabolic rate that was 1.5-fold (males) and 1.8-fold (females) lower than basal metabolic rate (BMR) measured in this and a previous study. The difference was probably attributable to birds being stressed while they were held in the metabolic chamber or wearing a mask. Thus, previous measurements of metabolic rate under basal conditions or for incubating wandering albatrosses are likely to be overestimates. Combining the relationship between HR and metabolic rate for both sexes, we estimate that wandering albatrosses expend 147 kJ kg–1 day–1 to incubate their eggs. In addition, the cost of incubation was assumed to vary because (i) HR was higher during the day than at night, and (ii) there was an effect of wind chill (\u3c0°C) on basal HR. The presence of humans in the vicinity of the nest or after a band control was shown to increase HR for extended periods (2–3 h), suggesting that energy expenditure was increased as a result of the disturbance. Lastly, males and females reacted differently to handling in terms of HR response: males reacted more strongly than females before handling, whereas females took longer to recover after being handled

The effects of spatially heterogeneous prey distributions on detection patterns in foraging seabirds

Many attempts to relate animal foraging patterns to landscape heterogeneity are focused on the analysis of foragers movements. Resource detection patterns in space and time are not commonly studied, yet they are tightly coupled to landscape properties and add relevant information on foraging behavior. By exploring simple foraging models in unpredictable environments we show that the distribution of intervals between detected prey (detection statistics)is mostly determined by the spatial structure of the prey field and essentially distinct from predator displacement statistics. Detections are expected to be Poissonian in uniform random environments for markedly different foraging movements (e.g. L\'evy and ballistic). This prediction is supported by data on the time intervals between diving events on short-range foraging seabirds such as the thick-billed murre ({\it Uria lomvia}). However, Poissonian detection statistics is not observed in long-range seabirds such as the wandering albatross ({\it Diomedea exulans}) due to the fractal nature of the prey field, covering a wide range of spatial scales. For this scenario, models of fractal prey fields induce non-Poissonian patterns of detection in good agreement with two albatross data sets. We find that the specific shape of the distribution of time intervals between prey detection is mainly driven by meso and submeso-scale landscape structures and depends little on the forager strategy or behavioral responses.Comment: Submitted first to PLoS-ONE on 26/9/2011. Final version published on 14/04/201

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

1. In a highly dynamic airspace, flying animals are predicted to adjust foraging behaviour to variable wind conditions to minimize movement costs. 2. 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. 3. 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. 4. 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. 5. 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. 6. 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