1Long distance migration can increase lifetime fitness, but can be costly, incurring in-2 creased energetic expenses and higher mortality risks. Stopover and other en route be-3 haviors allow animals to rest and replenish energy stores and avoid or mitigate other 4 hazards during migration. Some animals, such as soaring birds, can subsidize the ener-5 getic costs of migration by extracting energy from flowing air. However, it is unclear how 6 these energy sources affect or interact with behavioral processes and stopover in long-7 distance soaring migrants. To understand these behaviors and the effects of processes 8 that might enhance use of flight subsidies, we developed a flexible mechanistic model to 9 predict how flight subsidies drive migrant behavior and movement processes. The novel 10 modelling framework incorporated time-varying parameters informed by environmental 11 covariates to characterize a continuous range of behaviors during migration. This model 12 framework was fit to GPS satellite telemetry data collected from a large soaring and op-13 portunist foraging bird, the golden eagle (Aquila chrysaetos), during migration in western 14 North America. Fitted dynamic model parameters revealed a clear circadian rhythm in 15 eagle movement and behavior, which was directly related to thermal uplift. Behavioral 16 budgets were complex, however, with evidence for a joint migrating/foraging behavior, 17 resembling a slower paced fly-and-forage migration, which could facilitate efficient refu-18 eling while still ensuring migration progress. In previous work, ecological and foraging 19 conditions are normally considered to be the key aspects of stopover location quality, 20 but taxa that can tap energy sources from moving fluids to drive migratory locomotion, 21 such as the golden eagle, may pace migration based on both foraging opportunities and 22 available flight subsidies.
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Keywords
24Bayesian, correlated random walk, golden eagle, movement ecology, soaring flight 25 Long-distance migration can relax competition and permit use of seasonally available 27 resources, helping many animals maximize lifetime fitness (Newton, 2008; Avgar et al., 28 2014). Those benefits, however, come at substantial costs, including greater vulnerability 29 to predators, uncertain conditions, mechanical wear, elevated energy expenditure, and 30 time (Alerstam and Hedenström, 1998; Clark and Butler, 1999; Hedenström, 2008; New-31 ton, 2008; Avgar et al., 2014). As many migrant species cannot store sufficient energy for 32 nonstop, long-distance migration, stopover evolved as a behavior for strategically resting 33 and refueling en route (Gill, 2007).
34Migrant species are adapted for utilizing either soaring or flapping flight, and the 35 different flight modes can relate to stopover strategy (Hedenström, 1993; Gill, 2007). 36 Generally, soaring flight is favorable for larger birds and flapping flight for smaller birds, 37 though the partitioning of time for each flight mode during migration is dependent on the 38 relativ...