Adult conspecific density affects Janzen-Connell patterns by modulating the recruitment exclusion zones

Plant-soil negative feedback (NF) is a well-established phenomenon that, by preventing the dominance of a single species, allows species coexistence and promotes the maintenance of biodiversity. At community scale, localized NF may cause the formation of exclusion zones under adult conspecifics leading to Janzen-Connell (JC) distribution. In this study, we explore the connection between adult density, either conspecifics or heterospecifics, on the probability of occurrence of JC distributions. Using an individual-based modelling approach, we simulated the formation of exclusion zones due to the build-up of NF in proximity of conspecific adult plants and assessed the frequency of JC distribution in relation to conspecifics and heterospecifics density ranging from isolated trees to closed forest stands. We found that JC recruitment distribution is very common in the case of an isolated tree when NF was strong and capable to form an exclusion zone under the parent tree. At very low NF intensity, a prevalence of the decreasing pattern was observed because, under such conditions, the inhibitory effect due to the presence of the mother tree was unable to overcome the clustering effect of the seed dispersal kernel. However, if NF is strong the JC frequency suddenly decreases in stands with a continuous conspecific cover likely as a result of progressive expansion of the exclusion zone surrounding all trees in closed forest stands. Finally, our simulations showed that JC distribution should not be frequent in the case of rare species immersed in a matrix of heterospecific adults. Overall, the model shows that a plant suffering from strong NF in monospecific stands can rarely exhibit a recruitment pattern fitting the JC model. Such counterintuitive results would provide the means to reconcile the well-established NF framework with part the forest ecologists' community that is still skeptical towards the JC model.

Our model highlights the complex interconnection between NF intensity, stand density, and recruitment patterns explaining where and why the JC distribution occurs. Moreover, predicting the occurrence of JC in relation to stand density we clarify the relevance of this ecological phenomenon for future integration in plant community frameworks.