(Camila Gómez, Nicholas J. Bayly, D. Ryan Norris, Stuart A. Mackenzie, Kenneth V. Rosenberg, Philip D. Taylor, Keith A. Hobson & Carlos Daniel Cadena 13 June 2017)
Although migration is an adaptive behaviour in a wide range of animals1,2,3, it is also thought to impose significant costs on individuals4. Studies on various migratory birds5,6,7, mammals8 and fish9 provide evidence that mortality can be higher during migration than during stationary periods of the annual cycle. In addition, work on birds10, 11 and insects12 indicates that migrating individuals often undergo significant metabolic and behavioural adjustments to fulfil the high energetic demands of migration. Time spent and energy used during migration can also determine subsequent breeding success10, 12,13,14,15, emphasizing the high costs that individuals pay when migrating. Because migration is costly, migratory organisms are expected to maximize their fitness behaviourally via minimizing either the time spent, energy consumed, or the risks incurred during migratory journeys16, 17.
In terms of time, the highest cost of migration is generally thought to be experienced during stopovers rather than during periods of flight18, 19, and birds rely on the time spent at stopover sites to rest and refuel for the next leg of their journeys20. Optimal migration theory provides a framework to study stopover behaviour and its consequences by testing whether migrants are time- or energy-minimizers using data on fuelling rate, stopover duration, fuel loads and potential flight ranges17. Individuals attempting to minimize the overall time spent on migration are expected to maximize the amount of fuel they can acquire at each stopover in the shortest time possible. A key consequence of this strategy is that it maximizes the distance that can be flown between stopovers18, 21. Consequently, the fuel loads (amount of fat carried) of a time-minimizer should be tightly linked to local conditions at stopover sites as well as to the conditions expected ahead because these conditions influence fuelling rates18, 21. Furthermore, stopover durations in time-minimizers are expected to have been shaped by or to respond directly to experienced fuelling conditions17, 18. Larger departure fuel loads should allow for longer flights and a faster overall pace of migration because individuals acquiring sufficient fuel in the shortest time possible will need to make fewer stopovers and be able to take more direct routes to their destination, including being able to fly over physical barriers or large areas of unsuitable habitat such as deserts or oceans rather than circumventing these areas22.