Abstract

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.

Highlights

  • Optimal foraging theory predicts that foraging animals should adjust their behaviour to maximize both time and energy efficiency (Pyke, 1984; Stephens, Brown, & Ydenberg, 2008; Ydenberg, Welham, Schmid-Hempel, Schmid-Hempel, & Beauchamp, 1994)

  • 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

  • When comparing outputs of hidden Markov models (HMMs) to finer scale immersion data for birds at South Georgia, there was a good match for directed flight and rest, but during area-restricted search (ARS), birds undertook finer scale sequences of wet and dry activity likely associated with prey capture attempts, which were not apparent from the lower frequency GPS fixes (Table S3 in Appendix S2)

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Summary

| INTRODUCTION

Optimal foraging theory predicts that foraging animals should adjust their behaviour to maximize both time and energy efficiency (Pyke, 1984; Stephens, Brown, & Ydenberg, 2008; Ydenberg, Welham, Schmid-Hempel, Schmid-Hempel, & Beauchamp, 1994). Large soaring birds are well-adapted to exploit a dynamic airspace (Hedenström, 1993; Richardson, 2011; Shepard, Williamson, & Windsor, 2016), extracting kinetic energy from wind for soaring–gliding flight (Pennycuick, 1982, 1998) This substantially reduces time spent flapping, which is metabolically costly compared with gliding (Duriez et al, 2014; Weimerskirch, Guionnet, Martin, Shaffer, & Costa, 2000). Due to sex differences in flight performance (Shaffer et al, 2001), we predict that behavioural responses of males will be more strongly influenced by wind Due to their higher wing loading, males should both remain in directed gliding flight, and be more likely to take off in stronger winds than females (H3)

| MATERIALS AND METHODS
Findings
| DISCUSSION
| CONCLUSIONS
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