1 Introduction Environmental heterogeneity and temporal variability are fundamental drivers of ecosystem structure and function (Levin 1992). Spatial heterogeneity provides a mosaic of habitats, and resources that support biodiversity (MacIntosh and Tonkin 2024; Yang et al. 2025), facilitate species coexistence (Stien et al. 2014), and enhances ecosystem resilience (Oliver et al. 2015).