Pervasive and persistent effects of ant invasion and fragmentation on native ant assemblages.

Biological invasions are a leading cause of global change, yet their long-term effects remain hard to predict. Invasive species can remain abundant for long periods of time, or exhibit population crashes that allow native communities to recover. The abundance and impact of non-native species may also be closely tied to temporally variable habitat characteristics. We investigated the long-term effects of habitat fragmentation and invasion by the Argentine ant (Linepithema humile) by resurveying ants in 40 scrub habitat fragments in coastal southern California that were originally sampled 21 years ago. At a landscape scale, fragment area, but not fragment age or Argentine ant mean abundance, continued to explain variation in native ant species richness; the species-area relationship between the two sample years did not differ in terms of slope or intercept. At local scales, over the last 21 years we detected increases in the overall area invaded (+36.7 %, estimated as the proportion of occupied traps) and the relative abundance of the Argentine ant (+121.95 %, estimated as mean number of workers in pitfall traps). Argentine ant mean abundance also increased inwards from urban edges in 2017 compared to 1996. The greater level of penetration into fragments likely reduced native ant richness by eliminating refugia for native ants in fragments that did not contain sufficient interior area. At one fragment where we sampled eight times over the last 21 years, Argentine ant mean abundance increased over time while the diversity of native ground-foraging ants declined from 14 to 4 species. Notably, native species predicted to be particularly sensitive to the combined effect of invasion and habitat loss were not detected at any sites in our recent sampling, including the army ant genus Neivamyrmex. Conversely, two introduced ant species (Brachymyrmex patagonicus and Pheidole flavens) that were undetected in 1996 are now common and widespread at our sites. Our results indicate that behaviorally and numerically dominant invasive species can maintain high densities and suppress native diversity for extended periods.