Darwin's naturalization conundrum:an unsolved paradox in invasion ecology
In On The Origin of Species,Darwin proposed two seemingly contradictory hypotheses that predict how the phylogenetic relatedness of alien species to native species can either facilitate or impede their naturalization and invasion.These two apparently opposing hypotheses,now known as"Darwin's naturalization conundrum",has garnered increasing attention over the past three decades with the advancement of phylogenetic and trait-based analyses.However,recent studies have often produced mixed results,posing a major challenge in reconciling this conundrum.In this paper,we provide a comprehensive overview of the essence and significance of Darwin's naturalization conundrum,along with a review of the research history and recent advancements in the field.Specifically,we emphasize recent progress made in relation to spatial scale,invasion stage,ecological difference measures,and controlled experiments.Additionally,we propose key directions for future research,including the consideration of environmental gradients and human activities,analysis of native community structures and dynamics during invasion,utilization of appropriate statistical methods and null models,and extending the conundrum's concepts to novel fields such as disease ecology.Addressing this 160-year-old conundrum will not only contribute to the development and refinement of invasion ecology theory but also provide valuable scientific guidance for the prevention and management of biological invasions.