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Ecologists have achieved much progress in the study of mechanisms that maintain species coexistence and diversity. In this paper, we reviewed a wide range of past research related to these topics, focusing on five theoretical bodies: (1) coexistence by niche differentiation, (2) coexistence without niche differentiation, (3) coexistence along environmental stress gradients, (4) coexistence under non-equilibrium versus equilibrium conditions, and (5) modern perspectives. From the review, we identified that there are few models that can be generally and confidently applicable to different ecological systems. This problem arises mainly because most theories have not been substantiated by enough empirical research based on field data to test various coexistence hypotheses at different spatial scales. We also found that little is still known about the mechanisms of species coexistence under harsh environmental conditions. This is because most previous models treat disturbance as a key factor shaping community structure, but they do not explicitly deal with stressful systems with non-lethal conditions. We evaluated the mainstream ideas of niche differentiation and stochasticity for the coexistence of plant species across salt marsh creeks in southwestern Denmark. The results showed that diversity indices, such as Shannon–Wiener diversity, richness, and evenness, decreased with increasing surface elevation and increased with increasing niche overlap and niche breadth. The two niche parameters linearly decreased with increasing elevation. These findings imply a substantial influence of an equalizing mechanism that reduces differences in relative fitness among species in the highly stressful environments of the marsh. We propose that species evenness increases under very harsh conditions if the associated stress is not lethal. Finally, we present a conceptual model of patterns related to the level of environmental stress and niche characteristics along a microhabitat gradient (i.e., surface elevation). The ecology of stressful systems with non-lethal conditions will be increasingly important as ongoing global-scale climate change extends the period of chronic stresses that are not necessarily fatal to inhabiting plants. We recommend that more ecologists continue this line of research.

Coexistence of plant species under harsh environmental conditions: an evaluation of niche differentiation and stochasticity along salt marsh creeks