Functional relationships reveal keystone effects of the gopher tortoise on vertebrate diversity in a longleaf pine savanna

Keystone species are important drivers of diversity patterns in many ecosystems. Their effects on ecological processes are fundamental to understanding community dynamics, making them attractive conservation targets for ecosystem management. However, many studies assume keystone effects are constant. By developing functional relationships of species’ effects and assessing how they vary with context, we can design more efficient conservation strategies to maintain keystone impacts. The threatened gopher tortoise (Gopherus polyphemus) is presumed to be a keystone species promoting biodiversity in endangered longleaf pine ecosystems of the Southeastern Coastal Plain, USA. Although many commensals use tortoise burrows, their putative keystone influence on emergent diversity patterns lacks critical evaluation. We quantified the functional relationship between tortoise burrow density and non-volant vertebrate diversity in a longleaf pine savanna, located in central Florida. Tortoise burrow density had a positive effect on vertebrate diversity and evenness but did not affect species richness. This relationship was robust across fire disturbance regimes and was the primary factor explaining diversity at the local scale. Our results demonstrate keystone effects of the gopher tortoise through an ecosystem engineering mechanism. Continued gopher tortoise population declines will have large, negative impacts on vertebrate diversity in this biodiversity hotspot. Therefore, maintaining gopher tortoise populations is critical to effectively conserve dependent species and the function of endangered longleaf pine ecosystems. We show that developing a functional understanding of keystone relationships (not a binomial categorization) can lead to important insights into community processes.