Nurse abundance determines plant facilitation networks of subtropical forest–grassland ecotone

The theory of complex networks has been recently used to explain ecological associations between nurses and seedlings in plant facilitation systems. The structure of these networks is potentially affected by morphological, ecological and evolutionary factors that can determine the facilitative interactions. In the present study, we evaluate the role of the projected crown area of the nurse plant, the dispersal syndrome, the abundance and the phylogeny relationships with seedlings as drivers of network structure in facilitation networks. To test these parameters, we used an ecotonal forest–grassland system in southern Brazil that experienced historical forest expansion over the grassland in the last few centuries. In two State Parks, Guartelá (GUA) and Vila Velha (VVL), we sampled tree species (nurses and seedlings) along four transects parallel to the forest edge, with five sampling stations along each transect. Tree seedlings (height >0.3 m) were sampled below the nurse's crown (i.e. isolated woody plants, height >1.5 m) and over the grassland area. A total of 160 nurses and 358 seedlings were sampled. Seedling abundance and richness were greater below a nurse's crown than on open grassland. Nurse abundance was the best predictor of the observed interaction matrix. Probabilistic matrices based on abundance, abundance and phylogeny, and dominance were able to predict the observed nestedness values, and they were also closest to the observed connectance, although all of the probabilistic matrices have underestimated this property. Specialization was predicted by abundance and phylogeny, abundance and projected crown area drivers for VVL, and the abundance based models were closest to the predicted specialization for GUA. The result of a similar pattern in predictability between sites indicates that the influences of morphology and evolutionary and ecological processes over the facilitation interactions are equivalent on a regional scale. Woody plant abundance was a key factor for the facilitation networks, driving forest expansion along the subtropical forest–grassland ecotone.