Trade-off between early emergence and herbivore susceptibility mediates exotic success in an experimental California plant community.

Ecological trade-offs are fundamental to theory in community ecology; critical for understanding species coexistence in diverse plant communities, as well as the evolution of diverse life-history strategies. Invasions by exotic species can provide insights into the importance of trade-offs in community assembly, because the ecological strategies of invading species often differ from those present in the native species pool. Exotic annual species have invaded many Mediterranean-climate areas around the globe, and often germinate and emerge earlier in the growing season than native species. Early-season growth can enable exotic annual species to preempt space and resources, competitively suppressing later-emerging native species; however, early-emerging individuals may also be more apparent to herbivores. This suggests a potential trade-off between seasonal phenology and susceptibility to herbivory. To evaluate this hypothesis, we monitored the emergence and growth of 12 focal species (six each native and exotic) in monoculture and polyculture, while experimentally excluding generalist herbivores both early and later in the growing season. Consistent with past studies, the exotic species emerged earlier than native species. Regardless of species origin, earlier-emerging species achieved greater biomass by the end of the experiment, but were more negatively impacted by herbivory, particularly in the early part of the growing season. This greater impact of early-season herbivory on early-active species led to a reduction in the competitive advantage of exotic species growing in polyculture, and improved the performance of later-emerging natives. Such a trade-off between early growth and susceptibility to herbivores could be an important force in community assembly in seasonal herbaceous-dominated ecosystems. These results also show how herbivore exclusion favors early-active exotic species in this system, with important implications for management in many areas invaded by early-active exotic species.