Topographic heterogeneity triggers complementary cascades that enhance ecosystem multifunctionality

Topographic heterogeneity sets the stage for community assembly, but its effects on ecosystem functioning remain poorly understood. Here, we test the hypothesis that topographic heterogeneity underpins multiple cascading species interactions and functional pathways that indirectly control multifunctionality. To do so, we combined experimental manipulation of a form of topographic heterogeneity on rocky shores (holes of various sizes) with a comprehensive assessment of naturally assembled communities and multifunctionality. Structural equation modeling indicated that heterogeneity: (1) enhanced biodiversity by supporting filter feeder richness; (2) triggered a facilitation cascade via reef-forming (polychaete) and biomass-dominant (macroalga) foundation species, which in turn broadly supported functionally diverse epibiotic and understory assemblages; and (3) inhibited a key consumer (limpet). The model supported that these mechanisms exerted complementary positive effects on individual functions (e.g., water filtration, ecosystem metabolism, nutrient uptake) and, in turn, collectively enhanced multifunctionality. Topographic heterogeneity may therefore serve as a cornerstone physical attribute by initiating multiple cascades that propagate through ecological communities via foundation species, ultimately manifesting disproportionate effects on ecosystem multifunctionality.