Impacts of Invasive Species on Food Webs: A Review of Empirical Data

We review empirical studies on how bioinvasions alter food webs and how a food-web perspective may change their prediction and management. Predation is found to underlie the most spectacular damage in invaded systems, sometimes cascading down to primary producers. Indirect trophic effects (exploitative and apparent competition) also affect native species, but rarely provoke extinctions, while invaders often have positive bottom-up effects on higher trophic levels. As a result of these trophic interactions, and of nontrophic ones such as mutualisms or ecosystem engineering, invasions can profoundly modify the structure of the entire food web. While few studies have been undertaken at this scale, those that have highlight how network properties such as species richness, phenotypic diversity, and functional diversity, limit the likelihood and impacts of invasions by saturating niche space. Vulnerable communities have unsaturated niche space mainly because of evolutionary history in isolation (islands), dispersal limitation, or anthropogenic disturbance. Evolution also modulates the insertion of invaders into a food web. Exotics and natives are evolutionarily new to one another, and invasion tends to retain alien species that happen to have advantage over residents in trophic interactions. Resident species, therefore, often rapidly evolve traits to better tolerate or exploit invaders-a process that may eventually restore more balanced food webs and prevent extinctions. We discuss how network-based principles might guide management policies to better live with invaders, rather than to undertake the daunting (and often illusory) task of eradicating them one by one.