Secondary foundation species as drivers of trophic and functional diversity: evidence from a tree-epiphyte system.

Facilitation cascades arise where primary foundation species facilitate secondary (dependent) foundation species, and collectively, they increase habitat complexity and quality to enhance biodiversity. Whether such phenomena occur in nonmarine systems and if secondary foundation species enhance food web structure (e.g., support novel feeding guilds) and ecosystem function (e.g., provide nursery for juveniles) remain unclear. Here we report on field experiments designed to test whether trees improve epiphyte survival and epiphytes secondarily increase the number and diversity of adult and juvenile invertebrates in a potential live oak-Tillandsia usneoides (Spanish moss) facilitation cascade. Our results reveal that trees reduce physical stress to facilitate Tillandsia, which, in turn, reduces desiccation and predation stress to facilitate invertebrates. In experimental removals, invertebrate total density, juvenile density, species richness and H' diversity were 16, 60, 1.7, and 1.5 times higher, and feeding guild richness and H' were 5 and 11 times greater in Tillandsia-colonized relative to Tillandsia-removal limb plots. Tillandsia enhanced communities similarly in a survey across the southeastern United States. These findings reveal that a facilitation cascade organizes this widespread terrestrial assemblage and expand the role of secondary foundation species as drivers of trophic structure and ecosystem function. We conceptualize the relationship between foundation species' structural attributes and associated species abundance and composition in a Foundation Species-Biodiversity (FSB) model. Importantly, the FSB predicts that, where secondary foundation species form expansive and functionally distinct structures that increase habitat availability and complexity within primary foundation species, they generate and maintain hot spots of biodiversity and trophic interactions.