The effect of environmental heterogeneity on clonal behaviour of Prunella vulgaris L.

Melinis minutiflora Beauv. (Poaceae) is an African grass that is invading mid-elevation Trachypogon savannas in Venezuela. The objective of this study was to investigate the influence of soil fertility, competition and soil disturbance in facilitating Melinis' invasion and growth in these savanna sites. We manipulated soil fertility by adding nitrogen (+N), phosphorus and potassium (+PK), or nitrogen, phosphorus, and potassium (+NPK). We simultaneously manipulated the competitive environment by clipping background vegetation. In a separate experiment, we mechanically disrupted the soil to simulate disturbance. We hypothesized that germination and growth were bottlenecks to early establishment in undisturbed savanna, but that disturbance would alleviate those bottlenecks. We measured Melinis seed germination and subsequent establishment by adding seeds to all plots. We examined Melinis growth by measuring biomass of Melinis seedling transplants, 11 months after they were placed into treatment plots. Germination and establishment of Melinis from seed was extremely low. Of the 80,000 seeds applied in the experiment, only 28 plants survived the first growing season. Mortality of Melinis seedling transplants was lowest in PK fertilized plots, but in the absence of PK mortality increased with N additions and clipping. By contrast, fertilization of the savanna with NPK greatly increased Melinis seedling biomass and this effect was greatly enhanced when competition was reduced (e.g. clipping). Melinis transplant growth responded strongly to soil disturbance- a response not fully explained by removal of competitors (clipping) or changes in soil nutrients and moisture. We suspect that disruption of the soil structure allowed for greater root proliferation and subsequent plant growth. We believe that native savanna is relatively resistant to Melinis invasion, since Melinis seedlings persisted in intact savanna but exhibited little or no growth during the first year. The significant enhancement of Melinis seedling growth with clipping and nutrient additions suggests that low soil nutrients and the presence of native savanna species are important factors in the ability of native savanna to resist Melinis establishment. However, the potential for Melinis growth increases enormously with soil disturbance.