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Natl. Acad. Sci. U.S.A."],"published-print":{"date-parts":[[2026,6,16]]},"abstract":"<jats:p>The spread of invasive species poses a major threat to global biodiversity, yet predicting successful establishment of exotic species in novel environments remains challenging. Darwin\u2019s naturalization conundrum is a longstanding debate over whether exotic species closely or distantly related to native communities are more likely to succeed. Despite its long history, empirical studies continue to yield conflicting evidence. Here, we introduce the dark diversity concept, which refers to species that could theoretically inhabit a site but are currently absent. Combined with observed diversity, dark diversity integrates information on the potential diversity (site-specific species pool size) of the resident community and how completely that pool is locally present (community completeness). Analyzing a 340-y record of successful and failed fish introductions across 516 Swedish lakes, we showed that the effect of phylogenetic relatedness on invasion outcome depends on dark diversity context. Exotics closely related to resident species were more likely to establish in communities with smaller species pools and higher completeness, whereas phylogenetically distant exotic species were more successful in communities with larger species pools and lower completeness. Models with observed species richness obtained considerably less support. Thus, integrating the dark diversity framework clarifies the contrasting effects of phylogenetic relatedness on invasion outcomes, and helps reconcile this 160-y-old conundrum.<\/jats:p>","DOI":"10.1073\/pnas.2604929123","type":"journal-article","created":{"date-parts":[[2026,6,10]],"date-time":"2026-06-10T17:38:23Z","timestamp":1781113103000},"update-policy":"https:\/\/doi.org\/10.1073\/pnas.cm10313","source":"Crossref","is-referenced-by-count":1,"title":["Dark diversity framework reconciles Darwin\u2019s naturalization conundrum for freshwater fish invasions"],"prefix":"10.1073","volume":"123","author":[{"ORCID":"https:\/\/orcid.org\/0009-0002-2130-1067","authenticated-orcid":false,"given":"Wen-gang","family":"Zhang","sequence":"first","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02n96ep67","id-type":"ROR","asserted-by":"publisher"}],"name":"Research Center for Global Change and Ecological Forecasting, Zhejiang Tiantong Forest Ecosystem National Observation and Research Station, Institute of Eco-Chongming, School of Ecological and Environmental Sciences, East China Normal University","place":["Shanghai, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-2906-6471","authenticated-orcid":false,"given":"Meng","family":"Xu","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/034t30j35","id-type":"ROR","asserted-by":"publisher"}],"name":"Key Laboratory of Lake and Watershed Science for Water Security, Institute of Hydrobiology, Chinese Academy of Sciences","place":["Wuhan, China"]},{"id":[{"id":"https:\/\/ror.org\/04and7w57","id-type":"ROR","asserted-by":"publisher"}],"name":"Key Laboratory of Prevention and Control for Aquatic Invasive Alien Species, Ministry of Agriculture and Rural Affairs, Pearl River Fisheries Research Institute, Chinese Academy of Fishery Sciences","place":["Guangzhou, China"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3747-8572","authenticated-orcid":false,"given":"Shu-ya","family":"Fan","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/02n96ep67","id-type":"ROR","asserted-by":"publisher"}],"name":"Research Center for Global Change and Ecological Forecasting, Zhejiang Tiantong Forest Ecosystem National Observation and Research Station, Institute of Eco-Chongming, School of Ecological and Environmental Sciences, East China Normal University","place":["Shanghai, China"]},{"id":[{"id":"https:\/\/ror.org\/0546hnb39","id-type":"ROR","asserted-by":"publisher"}],"name":"Ecology, Department of Biology, University of Konstanz","place":["Konstanz, Germany"]},{"id":[{"id":"https:\/\/ror.org\/0415vcw02","id-type":"ROR","asserted-by":"publisher"}],"name":"The Forest Risk Research Centre, Faculty of Forestry and Wood Sciences, Czech University of Life Sciences Prague","place":["Prague, Czech Republic"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-5617-7524","authenticated-orcid":false,"given":"Jonathan A.","family":"Bennett","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/010x8gc63","id-type":"ROR","asserted-by":"publisher"}],"name":"Department of Plant Sciences, University of Saskatchewan","place":["Saskatoon, Canada"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2770-254X","authenticated-orcid":false,"given":"Ross N.","family":"Cuthbert","sequence":"additional","affiliation":[{"id":[{"id":"https:\/\/ror.org\/00hswnk62","id-type":"ROR","asserted-by":"publisher"}],"name":"Research Centre in Sustainable Energy, Institute for Global Food Security, School of Biological Sciences, Queen\u2019s University Belfast","place":["Belfast, United Kingdom"]}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Jaimie T. 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