The response of migratory populations to phenological change: a Migratory Flow Network modelling approach.

Declines in migratory species have been linked to anthropogenic climate change through phenological mismatch, which arises due to asynchronies between the timing of life-history events (such as migration) and the phenology of available resources. Long-distance migratory species may be particularly vulnerable to phenological change in their breeding ranges, since the timing of migration departure is based on environmental cues at distant non-breeding sites. Migrants may, however, be able to adjust migration speed en route to the breeding grounds, and thus, ability of migrants to update their timing of migration may depend critically on stopover frequency during migration; however, understanding how migratory strategy influences population dynamics is hindered by a lack of predictive models explicitly linking habitat quality to demography and movement patterns throughout the migratory cycle. Here, we present a novel modelling framework, the Migratory Flow Network (MFN), in which the seasonally varying attractiveness of breeding, winter and stopover regions drives the direction and timing of migration based on a simple general flux law. We use the MFN to investigate how populations respond to shifts in breeding site phenology based on their frequency of stopover and ability to detect and adapt to these changes. With perfect knowledge of advancing phenology, 'jump' migrants (low-frequency stopover) require more adaptation for populations to recover than 'hop' and 'skip' (high or medium frequency stopover) migrants. If adaptation depends on proximity, hop and skip migrants' populations can recover but jump migrants cannot adjust and decline severely. These results highlight the importance of understanding migratory strategies and maintaining high-quality stopover habitat to buffer migratory populations from climate-induced mismatch. We discuss how MFNs could be applied to diverse migratory taxa and highlight the potential of MFNs as a tool for exploring how migrants respond to other environmental changes such as habitat loss.