Abstract: Density dependence, both conspecific and heterospecific, is widely recognized as a crucial driver of plant species diversity. However, treating multiple heterospecific species as a homogeneous group obviously overlooks the variability in the impacts of different heterospecific neighbors on the survival or growth of focal species. In this study, we developed the Static-Dynamic Coupled Interspecific Association Classification Framework (SDIACF), which categorizes heterospecific neighbors based on their positive/negative interspecific associations and dynamic changes with focal species. We further used generalized linear mixed-effect models to analyze how conspecific and various heterospecific neighbors classified by the above framework influenced the survival and growth rates of the focal species. Our results revealed that heterospecific neighbors, deconstructed using SDIACF, exerted distinct effects on the focal species. Specifically, regardless of their initial interspecific association with the focal species, heterospecific neighbors with more negative associations showed a negative impact on the focal species, while those with more positive associations showed a positive effect. However, among heterospecific neighbors exhibiting identical dynamic interspecific associations, positively associated neighbors were slightly more conducive to the survival of the focal species than negatively associated ones, but slightly detrimental to its growth. In summary, our results demonstrated that heterospecific neighbors are not a homogeneous entity but play important and complex roles in species coexistence. The development of SDIACF not only constitutes a significant supplement to traditional density dependence research but also offers a novel perspective for further exploring species coexistence.

Key words: Static-Dynamic Coupled Interspecific Association, Classification of heterospecific neighbors, Heterospecific density dependence, Conspecific density dependence, Spatial distribution dynamics