Regional climate and local-scale biotic acceptance explain native-exotic richness relationships in Australian annual plant communities

Native and exotic species richness is expected to be negatively related at small spatial scales Where individuals interact, and positive at larger spatial scales as a greater variety of habitats are sampled. However, a range of native- exotic richness relationships (NERRs) have been reported, including positive at small scales and negative at larger scales. We present a hierarchical meta-community framework to explain how contrasting NERRs may emerge across scales and study systems, and then apply this framework to NERRs in an invaded winter annual plant system in southwest Western Australia. We analysed NERRs at increasing spatial scales from neighbourhoods (0.09 m(2)) to communities (225 m(2)) to metacommunities (greater than 10 ha) within a multilevel structural equation model. In contrast to many previous studies, native and exotic richness were positively related at the neighbourhood scale and were not significantly associated at larger scales. Heterogeneity in soil surface properties was weakly, but positively, associated with native and exotic richness at the community scale. Metacommunity exotic richness increased strongly with regional temperature and moisture availability, but relationships for native richness were negative and much weaker. Thus, we show that neutral NERRs can emerge at larger scales wing to differential climatic filtering of native and exotic species pools.