Anthropogenic disturbance and the spread of non-native species disrupt natural communities, but also create novel interactions between species. By-product mutualisms, in which benefits accrue as side effects of partner behaviour or morphology, are often non-specific and hence may persist in novel ecosystems. We tested this hypothesis for a two-way by-product mutualism between epiphytic ferns and their ant inhabitants in the Bornean rain forest, in which ants gain housing in root-masses while ferns gain protection from herbivores. Specifically, we assessed how the specificity (overlap between fern and ground-dwelling ants) and the benefits of this interaction are altered by selective logging and conversion to an oil palm plantation habitat. We found that despite the high turnover of ant species, ant protection against herbivores persisted in modified habitats. However, in ferns growing in the oil palm plantation, ant occupancy, abundance and species richness declined, potentially due to the harsher microclimate. The specificity of the fern-ant interactions was also lower in the oil palm plantation habitat than in the forest habitats. We found no correlations between colony size and fern size in modified habitats, and hence no evidence for partner fidelity feedbacks, in which ants are incentivised to protect fern hosts. Per species, non-native ant species in the oil palm plantation habitat (18 % of occurrences) were as important as native ones in terms of fern protection and contributed to an increase in ant abundance and species richness with fern size. We conclude that this by-product mutualism persists in logged forest and oil palm plantation habitats, with no detectable shift in partner benefits. Such persistence of generalist interactions in novel ecosystems may be important for driving ecosystem functioning.

Faculty of Science University of South Bohemia Branišovská 31 370 05 České Budějovice Czech Republic

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