Invader–invader mutualism influences land snail community composition and alters invasion success of alien species in tropical rainforest

Abstract

Mutualism between invaders may alter a key characteristic of the recipient community, leading to the entry or in situ release of other exotic species. We considered whether mutualism between invasive yellow crazy ant Anoplolepis gracilipes and exotic honeydew-producing scale insects indirectly facilitated land snails (exotic and native) via the removal of a native omnivore, the red land crab Gecarcoidea natalis. In plateau rainforest on Christmas Island, Indian Ocean, the land snail community was surveyed at 28 sites representing four forest states that differed in the density of red crabs, the abundance of yellow crazy ants and management history. One-way ANOVAs and multivariate analyses were used to determine differences in land snail species abundance and composition between forest states. Sample-based rarefaction was used to determine differences in species richness. The removal of the red land crab by supercolonies of yellow crazy ants was associated with a significant increase in the abundance of both invasive (14 species) and native (four species) land snails. Compositional differences in the land snail community were driven most strongly by the significantly greater abundance of a few common species in forest states devoid of red crabs. In forest where the crab population had recovered following management for ants, the land snail assemblage did not differ from intact, uninvaded forest. The land snail community was dominated by exotic species that can coexist alongside red crabs in rainforest uninvaded by exotic ants and scale insects. However, the ant–scale mutualism significantly increased land snail abundance and altered their composition indirectly though the alteration of the recipient community. We suggest these constitute ‘population-release’ secondary invasion in which the impacts of previously successful invaders facilitate a significant increase in abundance of other exotic species already established at low density within the community. Understanding facilitative interactions between invaders and indirect consequences of impacts will provide invaluable insights for conservation in heavily invaded ecosystems.