Linking animal migration and ecosystem processes: Data-driven simulation of propagule dispersal by migratory herbivores

Animal migration is a key process underlying active subsidies and species dispersal over long distances, which affects the connectivity and functioning of ecosystems. Despite much research describing patterns of where animals migrate, we still lack a framework for quantifying and predicting how animal migration affects ecosystem processes. In this study, we aim to integrate animal movement behavior and ecosystem functioning by developing a predictive modeling framework that can inform ecosystem management and conservation. We propose a framework to model individual-level migration trajectories between populations' seasonal ranges as well as the resulting dispersal and fate of propagules carried by the migratory animals, which can be calibrated using empirical data at every step of the modeling process. As a case study, we applied our framework to model the spread of guava seeds, Psidium guajava, by a population of migratory Galapagos tortoises, Chelonoidis porteri, across Santa Cruz Island. Galapagos tortoises are large herbivores that transport seeds and nutrients across the island, while Guava is one of the most problematic invasive species in the Galapagos archipelago. Our model can predict the pattern of spread of guava seeds alongside tortoises' downslope migration range, and it identified areas most likely to see establishment success. Our results show that Galapagos tortoises' seed dispersal may particularly contribute to guava range expansion on Santa Cruz Island, due to both long gut retention time and tortoise's long-distance migration across vegetation zones. In particular, we predict that tortoises are dispersing a significant amount of guava seeds into the Galapagos National Park, which has important consequences for the native flora. The flexibility and modularity of our framework allow for the integration of multiple data sources. It also allows for a wide range of applications to investigate how migratory animals affect ecosystem processes, including propagule dispersal but also other processes such as nutrient transport across ecosystems. Our framework is also a valuable tool for predicting how animal-mediated propagule dispersal can be affected by environmental change. These different applications can have important conservation implications for the management of ecosystems that include migratory animals.