Factors influencing dynamics of two invasive c4 grasses in seasonally dry hawaiian woodlands

The introduced C₄ bunchgrass, Schizachyrium condensatum, is abundant in unburned, seasonally dry woodlands on the island of Hawaii, where it promotes the spread of fire. After fire, it is partially replaced by Melinis minutiflora, another invasive C₄ grass. Seed bank surveys in unburned woodland showed that Melinis seed is present in locations without adult plants. Using a combination of germination tests and seedling outplant experiments, we tested the hypothesis that Melinis was unable to invade the unburned woodland because of nutrient and/or light limitation. We found that Melinis germination and seedling growth are depressed by the low light levels common under Schizachyrium in unburned woodland. Outplanted Melinis seedlings grew rapidly to flowering and persisted for several years in unburned woodland without nutrient additions, but only if Schizachyrium individuals were removed. Nutrients alone did not facilitate Melinis establishment. Competition between Melinis and Schizachyrium naturally occurs when individuals of both species emerge from the seed bank simultaneously, or when seedlings of one species emerge in sites already dominated by individuals of the other species. When both species are grown from seed, we found that Melinis consistently outcompetes Schizachyrium, regardless of light or nutrient treatments. When seeds of Melinis were added to pots with well‐established Schizachyrium (and vice versa), Melinis eventually invaded and overgrew adult Schizachyrium under high, but not low, nutrients. By contrast, Schizachyrium could not invade established Melinis pots regardless of nutrient level. A field experiment demonstrated that Schizachyrium individuals are suppressed by Melinis in burned sites through competition for both light and nutrients. Overall, Melinis is a dominant competitor over Schizachyrium once it becomes established, whether in a pot or in the field. We believe that the dominance of Schizachyrium, rather than Melinis, in the unburned woodland is the result of asymmetric competition due to the prior establishment of Schizachyrium in these sites. If Schizachyrium were not present, the unburned woodland could support dense stands of Melinis. Fire disrupts the priority effect of Schizachyrium and allows the dominant competitor (Melinis) to enter the system where it eventually replaces Schizachyrium through resource competition.