It is generally accepted that disturbances increase community invasibility. Yet the role of disturbance in plant invasions may be less predictable than often assumed, due to the influence of environmental stochasticity and interactions between disturbance regimes. We evaluated the single and interactive effects of prescribed burning (large-scale, infrequent event) and animal diggings (small-scale, frequent events) on the invasion success of Gleditsia triacanthos L. in a tussock grassland relict of the Inland Pampa, Argentina. Tree seedling emergence and survival were monitored over 4 years, after adjusting for propagule pressure through copious seed addition to all disturbance treatments. Burning altered community structure by suppressing tussock grasses and promoting exotic forbs, whereas simulated, armadillo-like diggings had little impact on herbaceous composition. Overall, seedling emergence rather than survival represented the main demographic bottleneck for tree invasion. Tree establishment success varied among seedling cohorts emerged in different climatic years. In a dry year, emergence was only slightly affected by disturbances. In contrast, for two consecutive wet years, initial burning and armadillo-like diggings exerted strong, antagonistic effects on tree recruitment. Whereas fire alone increased recruitment, the simulated burrowing regime prevented seedling emergence in both burned and unburned plots. The latter effect might be explained by reduced soil moisture, and increased seed burial and predation in excavated patches. Thus, the impact of a single, large-scale perturbation promoting woody plant invasion was overridden by a regime of small-scale, frequent disturbances. Our results show that grassland invasibility was contingent on inter-annual climatic variation as well as unexpected interactions between natural and anthropogenic disturbance agents.

Plants infected with vertically transmitted fungal endophytes carry their microbial symbionts with them during dispersal into new areas. Yet, whether seed-borne endophytes enhance the host plant's ability to overcome colonisation barriers and to regenerate within invaded sites remains poorly understood. We examined how symbiosis with asexual endophytic fungi (Neotyphodium) affected establishment and seed loss to predators in the invasive annual grass Lolium multiflorum (Italian ryegrass) across contrasting successional plots. Italian ryegrass seeds with high and low endophyte incidence were sown into three communities: a 1-year-old fallow field, a 15-year-old grassland, and a 24-year-old forest, which conformed to an old-field chronosequence in the eastern Inland Pampa, Argentina. We found that endophyte infection consistently increased host population recruitment and reproductive output. Endophyte presence also enhanced aerial biomass production of ryegrass in a low recruitment year but not in a high recruitment year, suggesting that symbiotic effects on growth performance are density dependent. Endophyte presence reduced seed removal by rodents, although differential predation may not account for the increased success of infected grass populations. Overall, there was no statistical evidence for an endophyte-by-site interaction, indicating that the fungal endosymbiont benefitted host establishment regardless of large differences in biotic and abiotic environment among communities. Our results imply that hereditary endophytes may increase the chances for host grass species to pass various ecological filters associated with invasion resistance across a broad range of successional habitats.