The imbalance of nature: revisiting a Darwinian framework for invasion biology

A major implication of natural selection is that species from different parts of the world will vary in their efficiency in converting resources into offspring for a given type of environment. This insight, articulated by Darwin, is usually overlooked in more recent studies of invasion biology that are often based on the more modern Eltonian perspective of imbalanced ecosystems. We formulate a renewed Darwinian framework for invasion biology, the evolutionary imbalance hypothesis (EIH), based only on the action of natural selection in historically isolated populations operating within a global network of repeated environments. This framework predicts that successful invaders are more likely to come from biotic regions of high genetic potential (with independent lineages of large population size), experiencing a given environment for many generations and under strong competition from other lineages.

Global.

We test the predictive power of this framework by examining disparities in recent species exchanges between global biotic regions, including patterns of plant invasions across temperate regions and exchanges of aquatic fauna as a result of modern canal building.

Our framework successfully predicts global invasion patterns using phylogenetic diversity of the world's biotic regions as a proxy that reflects their genetic potential, historical stability and competitive intensity, in line with the Darwinian expectation. Floristic regions of higher phylogenetic diversity are more likely to be source areas of invasive plants, and regions of lower phylogenetic diversity are more likely to be invaded. Similar patterns are evident for formerly isolated marine or freshwater assemblages that have been connected via canals.

We advocate an approach to understanding modern species invasions that recognizes the potential significance of both the original Darwinian explanation and the more modern view that emphasizes novel ecological or evolutionary mechanisms arising in the introduced range. Moreover, if biological invasions are a natural outcome of Darwinian evolution in an increasingly connected world, then invasive species should continue to displace native species and drive widespread shifts in the functioning of ecosystems.