Soil nitrogen availability favours the growth but not germination of secondary invaders after clearing invasive Acacia saligna

Invasive alien nitrogen-fixing species, such as Australian acacias, often leave a legacy of elevated available soil nitrogen after their removal. This legacy effect can facilitate secondary invasion by other alien species, thereby preventing natural restoration of areas being managed. To restore viable native plant communities in ecosystems where secondary invasion is a barrier to restoration, it is important to understand the soil legacy effects of invasions. Using Acacia saligna (Labill.) H. L. Wendl. (Fabaceae) invasions in the South African fynbos as case study, we determined (1) the extent to which levels of available soil nitrogen influence the germination and growth of secondary invaders; and (2) how this differs between secondary invader species. We chose five of the most common species that have been identified as secondary invaders after clearing invasive A. saligna in the Cape Flats Sand Fynbos: (i) Avena fatua (L.) (Poaceae), (ii) Briza maxima (L.) (Poaceae), (iii) Bromus diandrus (Roth.) (Poaceae), (iv) Hypochaeris radicata (L.) (Asteraceae), and (v) Raphanus raphanistrum (L.) (Brassicaceae). Using proportional fertigation, we created soil nitrate levels similar to those found in non-invaded areas (1mg/kg), and the lowest (3 mg/kg), median (7.5 mg/kg) and highest (12 mg/kg) levels typically found in areas previously invaded by A. saligna up to ten years after clearing. For each soil nitrate level, we germinated secondary invader seeds (five seeds per petri dish) in an incubator (five species × four soil nitrate levels × five replicates = 100 petri dishes). Furthermore, for each soil nitrate level, we grew secondary invaders (one plant per pot) in a greenhouse tunnel for five months (five species × four soil nitrate levels × five replicates = 100 pots). There was no significant relationship between germination success and soil nitrate level for any species. However, root and shoot dry mass were significantly positively correlated to soil nitrate level for all species. The relationship was nonetheless only linear for Bromus diandrus, Hypochaeris radicata, and Raphanus raphanistrum. These results indicate that the legacy of elevated available soil nitrogen does not have an effect on the germination of secondary invaders, but favours their growth. Therefore, secondary invasion can be managed through a reduction in soil nitrogen levels. However, secondary invader growth does not decrease at the same rate for all species in response to a decrease in available soil nitrogen. Thus, the use of soil nitrogen reduction as a tool for the management of secondary invasions should take into account the species being managed. However, given that multiple secondary invader species often dominate a restoration site, restoration efforts should reduce soil nitrogen to non-invaded levels to account for the different response rates to available soil nitrogen, and simply management efforts.