Stand age, degree of encroachment and soil characteristics modulate changes of C and N cycles in dry grassland soils invaded by the N2-fixing shrub Amorpha fruticosa

The N2-fixing shrub Amorpha fruticosa L. is rapidly spreading in the dry riparian natural grasslands of Europe, altering ecosystem functions and depleting plant diversity. Alteration of the N cycle represents the key factor involved in invasions by N2-fixing plants with cascading effects on plant species richness. We hypothesized that A. fruticosa encroachment strongly impacts not only the N but also the C cycle and that the magnitude of such alterations may be modulated by soil characteristics. To test these hypotheses, we selected four river floodplains in North East of Italy and compared natural uninvaded grasslands with half invaded and completely invaded sites, based on A. fruticosa stand characteristic and relevant leaf traits and on soil properties related to soil texture and to C and N cycles. Soil organic matter mineralisation, ammonification and nitrification rates were determined. Soil nitrification increased remarkably with plant invasion while ammonification was significantly higher only in half invaded sites. Soil organic matter mineralisation, microbial biomass C sustained per soil organic C unit and nitrification positively correlated with stand age, regardless to the stage of the encroachment. Mineralisation and nitrification increased with soil organic C and total N in uninvaded and completely invaded sites, but decreased in half invaded sites. At the half invasion stage, trends in nitrification and CO2 mineralisation were transitionally reverted and remediation may be facilitated by less pronounced changes in soil properties compared to completely invaded sites. Direct effects of plant invasion are modulated by the action of soil characteristics such as soil organic C and clay contents, with soils rich in organic C showing larger nitrification and mineralisation rates.