Multiple seed dispersal modes of an invasive plant species on coastal dunes

Beach evening-primrose (Oenothera drummondii subsp. drummondii) is a perennial herb native to the coastal dunes of the Gulf of Mexico and southeastern USA. During the last century, the species was unintentionally introduced into coastal dune systems around the world. The purpose of this study was to explore the means of dispersal of this invasive in the newly established populations on the coastal dunes of Spain and to determine if they contribute significantly to the invasion process. After reconstructing the history of its spread in Spain from available records, we tested whether dispersion by marine currents can be a mechanism of colonization at regional scale, and whether the presence of native vertebrate participate in the expansion process at local scale. Through laboratory experiments, we demonstrated that the seeds of O. drummondii can be dispersed by seawater, since 0.63% present both buoyancy in seawater and subsequent germination after being washed with fresh water. This capacity for dispersal by marine currents could explain both the expansion of the species along the southwestern coast of Spain and the maintenance of its populations following disturbances such as storms. We also demonstrated that Iberian Hares and European Rabbits can disperse the seeds by endozoochory; seeds recovered from faecal pellets of these species were capable of germination (73 and 63%, respectively). Oenothera drummondii has become an invader only in coastal dunes where these dispersers are present, and with no control actions to regulate their populations. In conclusion, O. drummondii is an alien species with varying dispersal methods, polychory. This helps maintain already established populations and disperses seeds at a regional level, depending on the incidence of storms and the presence or absence of rabbits and hares for efficient local seed dispersion. Thus, management actions that restrict marine and animal dispersal likely have the benefit of limiting the invasive pot