A sampling-theory framework for richness-uniqueness relationships

Ecological uniqueness, commonly quantified by local contribution to beta diversity (LCBD), is important for understanding and conserving biodiversity. Its widely documented negative relationship with species richness has led to the view that species-poor communities may be disproportionately important for regional diversity. Here, we derive neutral expectations for richness-uniqueness relationships and show that negative slopes can arise from neutral assembly rather than necessarily reflecting an ecological signal of uniqueness. Neutral simulations validate the analytical expectations across abundance- and incidence-based LCBD formulations. Applying the framework to the Barro Colorado Island tropical forest and North American Breeding Bird Survey, we show that accounting for neutral sampling expectations substantially changes the ranking and spatial distribution of ecologically unique sites. By separating neutral sampling expectations from excess uniqueness, our framework provides a principled basis for interpreting ecological uniqueness and informing biodiversity conservation.