Invasional meltdown: invader-invader mutualism facilitates a secondary invasion.

In multiply invaded ecosystems, introduced species should interact with each other as well as with native species. Invader-invader interactions may affect the success of further invaders by altering attributes of recipient communities and propagule pressure. The invasional meltdown hypothesis (IMH) posits that positive interactions among invaders initiate positive population-level feedback that intensifies impacts and promotes secondary invasions. IMH remains controversial: few studies show feedback between invaders that amplifies their effects, and none yet demonstrate facilitation of entry and spread of secondary invaders. Our results show that supercolonies of an alien ant, promoted by mutualism with introduced honeydew-secreting scale insects, permitted invasion by an exotic land snail on Christmas Island, Indian Ocean. Modeling of land snail spread over 750 sites across 135 km2 over seven years showed that the probability of land snail invasion was facilitated 253-fold in ant supercolonies but impeded in intact forest where predaceous native land crabs remained abundant. Land snail occurrence at neighboring sites, a measure of propagule pressure, also promoted land snail spread. Site comparisons and experiments revealed that ant supercolonies, by killing land crabs but not land snails, disrupted biotic resistance and provided enemy-free space. Predation pressure on land snails was lower (28.6%), survival 115 times longer, and abundance 20-fold greater in supercolonies than in intact forest. Whole-ecosystem suppression of supercolonies reversed the probability of land snail invasion by allowing recolonization of land crabs; land snails were much less likely (0.79%) to invade sites where supercolonies were suppressed than where they remained intact. Our results provide strong empirical evidence for IMH by demonstrating that mutualism between invaders reconfigures key interactions in the recipient community. This facilitates entry of secondary invaders and elevates propagule pressure, propagating their spread at the whole-ecosystem level. We show that identification and management of key facilitative interactions in invaded ecosystems can be used to reverse impacts and restore resistance to further invasions.

Invasive ants compete with and modify the trophic ecology of hermit crabs on tropical islands.

Invasive species can dramatically alter trophic interactions. Predation is the predominant trophic interaction generally considered to be responsible for ecological change after invasion. In contrast, how frequently competition from invasive species contributes to the decline of native species remains controversial. Here, we demonstrate how the trophic ecology of the remote atoll nation of Tokelau is changing due to competition between invasive ants (Anoplolepis gracilipes) and native terrestrial hermit crabs (Coenobita spp.) for carrion. A significant negative correlation was observed between A. gracilipes and hermit crab abundance. On islands with A. gracilipes, crabs were generally restricted to the periphery of invaded islands. Very few hermit crabs were found in central areas of these islands where A. gracilipes abundances were highest. Ant exclusion experiments demonstrated that changes in the abundance and distribution of hermit crabs on Tokelau are a result of competition. The ants did not kill the hermit crabs. Rather, when highly abundant, A. gracilipes attacked crabs by spraying acid and drove crabs away from carrion resources. Analysis of naturally occurring N and C isotopes suggests that the ants are effectively lowering the trophic level of crabs. According to delta(15) N values, hermit crabs have a relatively high trophic level on islands where A. gracilipes have not invaded. In contrast, where these ants have invaded we observed a significant decrease in delta(15) N for all crab species. This result concurs with our experiment in suggesting long-term exclusion from carrion resources, driving co-occurring crabs towards a more herbivorous diet. Changes in hermit crab abundance or distribution may have major ramifications for the stability of plant communities. Because A. gracilipes have invaded many tropical islands where the predominant scavengers are hermit crabs, we consider that their competitive effects are likely to be more prominent in structuring communities than predation.

Competitive assembly of South Pacific invasive ant communities.

BACKGROUND: The relative importance of chance and determinism in structuring ecological communities has been debated for nearly a century. Evidence for determinism or assembly rules is often evaluated with null models that randomize the occurrence of species in particular locales. However, analyses of the presence or absence of species ignores the potential influence of species abundances, which have long been considered of major importance on community structure. Here, we test for community assembly rules in ant communities on small islands of the Tokelau archipelago using both presence-absence and abundance data. We conducted three sets of analyses on two spatial scales using three years of sampling data from 39 plots on 11 islands. RESULTS: First, traditional null model tests showed support for negative species co-occurrence patterns among plots within islands, but not among islands. A plausible explanation for this result is that analyses at larger spatial scales merge heterogeneous habitats that have considerable effects on species occurrences. Second, analyses of ant abundances showed that samples with high ant abundances had fewer species than expected by chance, both within and among islands. One ant species, the invasive yellow crazy ant Anoplolepis gracilipes, appeared to have a particularly strong effect on community structure correlated with its abundance. Third, abundances of most ant species were inversely correlated with the abundances of all other ants at both spatial scales. This result is consistent with competition theory, which predicts species distributions are affected by diffuse competition with suites of co-occurring species. CONCLUSION: Our results support a pluralistic explanation for ant species abundances and assembly. Both stochastic and deterministic processes interact to determine ant community assembly, though abundance patterns clearly drive the deterministic patterns in this community. These deterministic patterns were observed at two spatial scales. Results indicate that abundance-based null models may be more sensitive in detecting non-random patterns in community assembly than species co-occurrences analyses.