Habitat heterogeneity, environmental feedbacks, and species coexistence across timescales (code and figures)

Classic ecological theory explains species coexistence in variable
environments. While spatial variation is often treated as an intrinsic
feature of a landscape, it may be shaped and even generated by the
resident community. All species modify their local environment to some
extent, driving changes that can feed back to affect the composition and
coexistence of the community, potentially over timescales very different
from population dynamics. We introduce a simple, nested modeling framework
for community dynamics in heterogeneous environments, including the
possible evolution of heterogeneity over time due to community-environment
feedbacks. We use this model to derive analytical conditions for species
coexistence in environments where heterogeneity is either fixed or shaped
by feedbacks. Among other results, our approach reveals how dispersal and
environmental specialization interact to shape realized patterns of
habitat association and demonstrates that environmental feedbacks can tune
landscape conditions to allow the stable coexistence of any number of
species. Our flexible modeling framework helps explain feedback dynamics
that arise in a wide range of ecosystems and offers a generic platform for
exploring the interplay between species and landscape diversity.