Riverscape heterogeneity shapes population diversity for a migratory fish

Habitat patch dynamics can scale up to influence population demography and diversity with implications for resilience to environmental stochasticity. But how the spatial arrangement and size of habitat patches interact with other components of habitat heterogeneity to shape population diversity at larger spatial scales is not well understood. For riverine fishes, there is increasing evidence that tributary streams provide critical demographic support to main stem rivers. However, the extent to which main stem rivers rely on demographic contributions from tributaries, and the factors underlying this dependence, have not been assessed. Here, we used genetic stock identification to evaluate the effect of tributaries on population diversity of Yellowstone cutthroat trout (Oncorhynchus virginalis bouvieri) occupying the main stem Snake River, Wyoming, USA. We found that the main stem relied almost entirely on tributaries for demographic support, but main stem composition varied spatially among river sections. Distance between habitat patches, catchment area, and groundwater availability acted in concert to determine the contribution of specific tributaries to the main stem, but contributions were ultimately modulated by habitat connectivity. We also found evidence for multi-scale spatial structure in tributary contributions, providing insight into untested drivers of main stem river population diversity. Our results demonstrate how spatially discrete and distributed riverscape attributes influence population diversity at broader spatial scales, illustrating how ecosystem resilience emerges from the dynamic, two-way exchange of individuals and energy across habitat networks. Management plans for large rivers that address the ecological contributions of tributaries may be needed to achieve optimal outcomes. Similarly, conservation strategies that exclusively focus on headwater streams may fail to capture the broader habitat requirements necessary to maintain robust cold-water fish populations and associated recreational fisheries, particularly under global environmental change.