Invasiveness and impact of the non-native shrub Baccharis halimifolia in sea rush marshes: fine-scale stress heterogeneity matters

The invasion of the exotic dioecious shrub Baccharis halimifolia is transforming the estuarine communities of Southern Europe. Large scale gradients of salinity and flooding regime determine B. halimifolia zonations in salt marshes where the subhalophilous sea rush communities are the most affected by invasion. In this study we aim to (1) assess the invasion level and influence of B. halimifolia on native flora and to (2) quantify the performance of the exotic shrub in rush communities across fine-scale salinity and waterlogging gradients. Using floristic data collected in estuaries in Northern Spain we identified 3 sea rush community subtypes: low, medium and high salinity communities. B. halimifolia cover decreased from low to high salinity communities. Native species cover, richness and diversity and herbaceous-subshrub layer cover was significantly lower in invaded rush communities than in uninvaded ones. The reduction of the singular native estuarine species cover and richness was higher in the high and medium salinity community than in the low salinity community. Growth and reproductive traits measured on two consecutive years in invaded rush communities in Urdaibai Biosphere Reserve indicated that increased edaphic stress reduced B. halimifolia individual performance and enhanced attack by natural enemies. Moreover, leaf drop was more responsive to salinity in female than in male individuals. We conclude that fine-scale variations on edaphic stress played an important role in the invasibility of rush communities by affecting the individual performance of B. halimifolia, and might generate sex specific responses. The implications for rush marsh conservation are discussed.