{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,11,12]],"date-time":"2026-11-12T18:24:58Z","timestamp":1794507898303,"version":"4.9.1"},"reference-count":92,"publisher":"Springer Science and Business Media LLC","issue":"8","license":[{"start":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T00:00:00Z","timestamp":1783296000000},"content-version":"tdm","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"},{"start":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T00:00:00Z","timestamp":1783296000000},"content-version":"vor","delay-in-days":0,"URL":"https:\/\/www.springernature.com\/gp\/researchers\/text-and-data-mining"}],"content-domain":{"domain":["link.springer.com"],"crossmark-restriction":false},"short-container-title":["Nat Ecol Evol"],"DOI":"10.1038\/s41559-026-03115-0","type":"journal-article","created":{"date-parts":[[2026,7,6]],"date-time":"2026-07-06T09:01:39Z","timestamp":1783328499000},"page":"1419-1433","update-policy":"https:\/\/doi.org\/10.1007\/springer_crossmark_policy","source":"Crossref","is-referenced-by-count":7,"title":["A catastrophic marine mortality event caused by a complex algal bloom including the brevetoxin producer Karenia cristata"],"prefix":"10.1038","volume":"10","author":[{"ORCID":"https:\/\/orcid.org\/0000-0001-7096-1307","authenticated-orcid":false,"given":"Shauna A.","family":"Murray","sequence":"first","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-9047-2391","authenticated-orcid":false,"given":"Christopher J. S.","family":"Bolch","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Steve","family":"Brett","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-3729-8176","authenticated-orcid":false,"given":"Cheong Xin","family":"Chan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Mark","family":"Doubell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-0890-7953","authenticated-orcid":false,"given":"Hazel","family":"Farrell","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2219-8145","authenticated-orcid":false,"given":"Greta","family":"Gaiani","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-1905-5349","authenticated-orcid":false,"given":"Hannah","family":"Greenhough","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-8464-7343","authenticated-orcid":false,"given":"Gustaaf","family":"Hallegraeff","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Tamsyne","family":"Smith-Harding","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-7222-5979","authenticated-orcid":false,"given":"D. Tim","family":"Harwood","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0009-0006-0613-5151","authenticated-orcid":false,"given":"Robert G.","family":"Hatfield","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Neil","family":"MacDonald","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Ian","family":"Moody","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0002-0267-6368","authenticated-orcid":false,"given":"Hugo","family":"Bastos de Oliveira","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Lesley","family":"Rhodes","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-2951-2969","authenticated-orcid":false,"given":"Anne","family":"Rolton","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Andrew I.","family":"Selwood","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Justin R.","family":"Seymour","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0001-6082-0949","authenticated-orcid":false,"given":"Nachshon","family":"Siboni","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-1521-0601","authenticated-orcid":false,"given":"Anastasiia","family":"Snigirova","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Nikola","family":"Streiber","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-4415-782X","authenticated-orcid":false,"given":"Craig","family":"Styan","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Clinton","family":"Wilkinson","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]},{"ORCID":"https:\/\/orcid.org\/0000-0003-3241-4147","authenticated-orcid":false,"given":"Kirsty","family":"Smith","sequence":"additional","affiliation":[],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"297","published-online":{"date-parts":[[2026,7,6]]},"reference":[{"key":"3115_CR1","unstructured":"GlobalHAB. Fish-Killing Marine Algal Blooms: Causative Organisms, Ichthyotoxic Mechanisms, Impacts and Mitigation (eds Hallegraeff, G. M. et al.) (UNESCO-IOC\/SCOR, 2023)."},{"key":"3115_CR2","unstructured":"SA Marine Mortality Events 2025 - 2026. iNaturalist https:\/\/www.inaturalist.org\/projects\/sa-marine-mortality-events-2025-2026 (2025)."},{"key":"3115_CR3","unstructured":"Environment and Communications References Committee. in Algal Blooms in South Australia 139\u2013148 (Commonwealth of Australia, 2025); https:\/\/parlinfo.aph.gov.au\/parlInfo\/download\/committees\/reportsen\/RB000648\/toc_pdf\/AlgalbloomsinSouthAustralia.pdf"},{"key":"3115_CR4","doi-asserted-by":"publisher","first-page":"117","DOI":"10.1038\/s43247-021-00178-8","volume":"2","author":"GM Hallegraeff","year":"2021","unstructured":"Hallegraeff, G. M. et al. Perceived global increase in algal blooms is attributable to intensified monitoring and emerging bloom impacts. Commun. Earth Environ. 2, 117 (2021).","journal-title":"Commun. Earth Environ."},{"key":"3115_CR5","doi-asserted-by":"crossref","unstructured":"Murray, S. A. & Gaiani, G. Mystery solved: our tests reveal the tiny algae killing fish and harming surfers on SA beaches. The Conversation https:\/\/theconversation.com\/mystery-solved-our-tests-reveal-the-tiny-algae-killing-fish-and-harming-surfers-on-sa-beaches-252810 (2025).","DOI":"10.64628\/AA.w57gnk6af"},{"key":"3115_CR6","doi-asserted-by":"publisher","first-page":"358","DOI":"10.1086\/335488","volume":"109","author":"CC Davis","year":"1948","unstructured":"Davis, C. C. Gymnodinium brevis sp. nov., a cause of discolored water and animal mortality in the Gulf of Mexico. Bot. Gaz. 109, 358\u2013360 (1948).","journal-title":"Bot. Gaz."},{"key":"3115_CR7","doi-asserted-by":"publisher","first-page":"309","DOI":"10.2307\/1948575","volume":"18","author":"G Gunter","year":"1948","unstructured":"Gunter, G., Williams, R. H., Davis, C. C. & Smith, F. W. Catastrophic mass mortality of marine animals and coincident phytoplankton bloom on the west coast of Florida, November 1946 to August 1947. Ecol. Monogr. 18, 309\u2013324 (1948).","journal-title":"Ecol. Monogr."},{"key":"3115_CR8","doi-asserted-by":"publisher","first-page":"755","DOI":"10.1038\/nature435755a","volume":"435","author":"LJ Flewelling","year":"2005","unstructured":"Flewelling, L. J. et al. Red tides and marine mammal mortalities. Nature 435, 755\u2013756 (2005).","journal-title":"Nature"},{"key":"3115_CR9","doi-asserted-by":"publisher","first-page":"598","DOI":"10.1016\/j.hal.2008.11.010","volume":"8","author":"JH Landsberg","year":"2009","unstructured":"Landsberg, J. H., Flewelling, L. J. & Naar, J. Karenia brevis red tides, brevetoxins in the food web, and impacts on natural resources: decadal advancements. Harmful Algae 8, 598\u2013607 (2009).","journal-title":"Harmful Algae"},{"key":"3115_CR10","doi-asserted-by":"publisher","first-page":"707","DOI":"10.1016\/j.toxicon.2007.06.005","volume":"50","author":"JP Naar","year":"2007","unstructured":"Naar, J. P. et al. Brevetoxins, like ciguatoxins, are potent ichthyotoxic neurotoxins that accumulate in fish. Toxicon 50, 707\u2013723 (2007).","journal-title":"Toxicon"},{"key":"3115_CR11","doi-asserted-by":"publisher","first-page":"618","DOI":"10.1289\/ehp.7501","volume":"113","author":"LE Fleming","year":"2005","unstructured":"Fleming, L. E., Backer, L. C. & Baden, D. G. Overview of aerosolized Florida red tide toxins: exposures and effects. Environ. Health Perspect. 113, 618\u2013620 (2005).","journal-title":"Environ. Health Perspect."},{"key":"3115_CR12","doi-asserted-by":"publisher","first-page":"224","DOI":"10.1016\/j.hal.2010.08.006","volume":"10","author":"LE Fleming","year":"2011","unstructured":"Fleming, L. E. et al. Review of Florida red tide and human health effects. Harmful Algae 10, 224\u2013233 (2011).","journal-title":"Harmful Algae"},{"key":"3115_CR13","doi-asserted-by":"publisher","first-page":"1807","DOI":"10.1096\/fasebj.3.7.2565840","volume":"3","author":"DG Baden","year":"1989","unstructured":"Baden, D. G. Brevetoxins: unique polyether dinoflagellate toxins. FASEB J. 3, 1807\u20131817 (1989).","journal-title":"FASEB J."},{"key":"3115_CR14","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0260755","volume":"17","author":"RP Stumpf","year":"2022","unstructured":"Stumpf, R. P. et al. Quantifying Karenia brevis bloom severity and respiratory irritation impact along the shoreline of Southwest Florida. PLoS ONE 17, e0260755 (2022).","journal-title":"PLoS ONE"},{"key":"3115_CR15","doi-asserted-by":"publisher","first-page":"156","DOI":"10.1016\/j.hal.2011.10.020","volume":"14","author":"LE Brand","year":"2012","unstructured":"Brand, L. E., Campbell, L. & Bresnan, E. Karenia: the biology and ecology of a toxic genus. Harmful Algae 14, 156\u2013178 (2012).","journal-title":"Harmful Algae"},{"key":"3115_CR16","doi-asserted-by":"publisher","first-page":"163","DOI":"10.1016\/S1568-9883(03)00026-X","volume":"2","author":"HA Maga\u00f1a","year":"2003","unstructured":"Maga\u00f1a, H. A., Contreras, C. & Villareal, T. A. A historical assessment of Karenia brevis in the western Gulf of Mexico. Harmful Algae 2, 163\u2013171 (2003).","journal-title":"Harmful Algae"},{"key":"3115_CR17","doi-asserted-by":"publisher","first-page":"155","DOI":"10.1515\/bot-2024-0083","volume":"68","author":"M Al-Kandari","year":"2025","unstructured":"Al-Kandari, M. et al. Morphological and molecular characterization of Kareniaceae (Dinophyceae, Gymnodiniales) in Kuwait\u2019s waters. Bot. Mar. 68, 155\u2013173 (2025).","journal-title":"Bot. Mar."},{"key":"3115_CR18","doi-asserted-by":"publisher","first-page":"102645","DOI":"10.1016\/j.hal.2024.102645","volume":"137","author":"J Cen","year":"2024","unstructured":"Cen, J. et al. Five Karenia species along the Chinese coast: with the description of a new species, Karenia hui sp. nov. (Kareniaceae, Dinophyta). Harmful Algae 137, 102645 (2024).","journal-title":"Harmful Algae"},{"key":"3115_CR19","doi-asserted-by":"publisher","DOI":"10.1016\/j.hal.2022.102204","volume":"114","author":"M Iwataki","year":"2022","unstructured":"Iwataki, M. et al. Morphological variation and phylogeny of Karenia selliformis (Gymnodiniales, Dinophyceae) in an intensive cold-water algal bloom in eastern Hokkaido, Japan. Harmful Algae 114, 102204 (2022).","journal-title":"Harmful Algae"},{"key":"3115_CR20","first-page":"269","volume":"133","author":"KA Steidinger","year":"2008","unstructured":"Steidinger, K. A., Wolny, J. L. & Haywood, A. J. Identification of Kareniaceae (Dinophyceae) in the Gulf of Mexico. Nova Hedwig. Beih. 133, 269 (2008).","journal-title":"Nova Hedwig. Beih."},{"key":"3115_CR21","doi-asserted-by":"publisher","DOI":"10.3389\/fmars.2025.1582234","volume":"12","author":"Y Xu","year":"2025","unstructured":"Xu, Y. et al. The first recorded fish-killing bloom in the Beibu Gulf, China: caused by dinoflagellate Karenia selliformis. Front. Mar. Sci. 12, 1582234 (2025).","journal-title":"Front. Mar. Sci."},{"key":"3115_CR22","doi-asserted-by":"publisher","first-page":"47","DOI":"10.1016\/j.hal.2011.10.002","volume":"13","author":"PT Holland","year":"2012","unstructured":"Holland, P. T. et al. Novel toxins produced by the dinoflagellate Karenia brevisulcata. Harmful Algae 13, 47\u201357 (2012).","journal-title":"Harmful Algae"},{"key":"3115_CR23","doi-asserted-by":"publisher","first-page":"375","DOI":"10.2989\/18142320609504181","volume":"28","author":"P McNabb","year":"2006","unstructured":"McNabb, P., Rhodes, L., Adamson, J. & Holland, P. Brevetoxin\u2014an elusive toxin in New Zealand waters. Afr. J. Mar. Sci. 28, 375\u2013377 (2006).","journal-title":"Afr. J. Mar. Sci."},{"key":"3115_CR24","doi-asserted-by":"publisher","first-page":"393","DOI":"10.3390\/md19070393","volume":"19","author":"Z Amzil","year":"2021","unstructured":"Amzil, Z. et al. Monitoring the emergence of algal toxins in shellfish: first report on detection of brevetoxins in French Mediterranean mussels. Mar. Drugs 19, 393 (2021).","journal-title":"Mar. Drugs"},{"key":"3115_CR25","doi-asserted-by":"publisher","first-page":"85","DOI":"10.1016\/j.toxicon.2015.05.007","volume":"101","author":"N Fowler","year":"2015","unstructured":"Fowler, N., Tomas, C., Baden, D., Campbell, L. & Bourdelais, A. Chemical analysis of Karenia papilionacea. Toxicon 101, 85\u201391 (2015).","journal-title":"Toxicon"},{"key":"3115_CR26","doi-asserted-by":"publisher","first-page":"624","DOI":"10.2216\/i0031-8884-43-5-624.1","volume":"43","author":"MF De Salas","year":"2004","unstructured":"De Salas, M. F., Bolch, C. J. & Hallegraeff, G. M. Karenia asterichroma sp. nov. (Gymnodiniales, Dinophyceae), a new dinoflagellate species associated with finfish aquaculture mortalities in Tasmania, Australia. Phycologia 43, 624\u2013631 (2004).","journal-title":"Phycologia"},{"key":"3115_CR27","doi-asserted-by":"publisher","DOI":"10.1016\/j.hal.2022.102337","volume":"120","author":"TY Orlova","year":"2022","unstructured":"Orlova, T. Y. et al. A massive bloom of Karenia species (Dinophyceae) off the Kamchatka coast, Russia, in the fall of 2020. Harmful Algae 120, 102337 (2022).","journal-title":"Harmful Algae"},{"key":"3115_CR28","doi-asserted-by":"publisher","first-page":"5829","DOI":"10.1016\/S0040-4039(02)01171-1","volume":"43","author":"M Satake","year":"2002","unstructured":"Satake, M. et al. Gymnocin-A, a cytotoxic polyether from the notorious red tide dinoflagellate, Gymnodinium mikimotoi. Tetrahedron Lett. 43, 5829\u20135832 (2002).","journal-title":"Tetrahedron Lett."},{"key":"3115_CR29","doi-asserted-by":"publisher","first-page":"7093","DOI":"10.1016\/0040-4039(95)01434-J","volume":"36","author":"T Seki","year":"1995","unstructured":"Seki, T., Satake, M., Mackenzie, L., Kaspar, H. F. & Yasumoto, T. Gymnodimine, a new marine toxin of unprecedented structure isolated from New Zealand oysters and the dinoflagellate, Gymnodinium sp. Tetrahedron Lett. 36, 7093\u20137096 (1995).","journal-title":"Tetrahedron Lett."},{"key":"3115_CR30","doi-asserted-by":"publisher","first-page":"19","DOI":"10.1016\/j.toxicon.2014.03.004","volume":"84","author":"DT Harwood","year":"2014","unstructured":"Harwood, D. T., Shi, F., Satake, M. & Holland, P. T. A sensitive LC-MS\/MS assay for brevisulcenal and brevisulcatic acid toxins produced by the dinoflagellate Karenia brevisulcata. Toxicon 84, 19\u201327 (2014).","journal-title":"Toxicon"},{"key":"3115_CR31","doi-asserted-by":"publisher","first-page":"989","DOI":"10.1021\/jo802183n","volume":"74","author":"M Satake","year":"2009","unstructured":"Satake, M. et al. Brevisin: an aberrant polycyclic ether structure from the dinoflagellate Karenia brevis and its implications for polyether assembly. J. Org. Chem. 74, 989\u2013994 (2009).","journal-title":"J. Org. Chem."},{"key":"3115_CR32","doi-asserted-by":"publisher","first-page":"3465","DOI":"10.1021\/ol801243n","volume":"10","author":"M Satake","year":"2008","unstructured":"Satake, M., Bourdelais, A. J., Van Wagoner, R. M., Baden, D. G. & Wright, J. L. Brevisamide: an unprecedented monocyclic ether alkaloid from the dinoflagellate Karenia brevis that provides a potential model for ladder-frame initiation. Org. Lett. 10, 3465\u20133468 (2008).","journal-title":"Org. Lett."},{"key":"3115_CR33","doi-asserted-by":"publisher","first-page":"536","DOI":"10.1021\/np900541w","volume":"73","author":"LT Truxal","year":"2010","unstructured":"Truxal, L. T., Bourdelais, A. J., Jacocks, H., Abraham, W. M. & Baden, D. G. Characterization of tamulamides A and B, polyethers isolated from the marine dinoflagellate Karenia brevis. J. Nat. Prod. 73, 536\u2013540 (2010).","journal-title":"J. Nat. Prod."},{"key":"3115_CR34","doi-asserted-by":"publisher","first-page":"138","DOI":"10.1038\/s41579-023-00975-2","volume":"22","author":"C Kuhlisch","year":"2024","unstructured":"Kuhlisch, C., Shemi, A., Barak-Gavish, N., Schatz, D. & Vardi, A. Algal blooms in the ocean: hot spots for chemically mediated microbial interactions. Nat. Rev. Microbiol. 22, 138\u2013154 (2024).","journal-title":"Nat. Rev. Microbiol."},{"key":"3115_CR35","doi-asserted-by":"publisher","first-page":"e0133549","DOI":"10.1371\/journal.pone.0133549","volume":"10","author":"JJ Dorantes-Aranda","year":"2015","unstructured":"Dorantes-Aranda, J. J., Seger, A., Mardones, J. I., Nichols, P. D. & Hallegraeff, G. M. Progress in understanding algal bloom-mediated fish kills: the role of superoxide radicals, phycotoxins and fatty acids. PLoS ONE 10, e0133549 (2015).","journal-title":"PLoS ONE"},{"key":"3115_CR36","doi-asserted-by":"publisher","first-page":"557","DOI":"10.1016\/j.hal.2010.04.005","volume":"9","author":"BD Mooney","year":"2010","unstructured":"Mooney, B. D., Hallegraeff, G. M. & Place, A. R. Ichthyotoxicity of four species of gymnodinioid dinoflagellates (Kareniaceae, Dinophyta) and purified karlotoxins to larval sheepshead minnow. Harmful Algae 9, 557\u2013562 (2010).","journal-title":"Harmful Algae"},{"key":"3115_CR37","doi-asserted-by":"publisher","DOI":"10.1016\/j.jenvman.2024.123465","volume":"373","author":"Q Zhang","year":"2025","unstructured":"Zhang, Q. et al. Genetic diversity and distribution of Karenia in the eastern coastal seas of China and implications for the trends in Karenia blooms under global environmental changes. J. Environ. Manag. 373, 123465 (2025).","journal-title":"J. Environ. Manag."},{"key":"3115_CR38","doi-asserted-by":"publisher","first-page":"573","DOI":"10.1016\/j.hal.2008.11.002","volume":"8","author":"GA Vargo","year":"2009","unstructured":"Vargo, G. A. A brief summary of the physiology and ecology of Karenia brevis Davis red tides on the West Florida Shelf and of hypotheses posed for their initiation, growth, maintenance, and termination. Harmful Algae 8, 573\u2013584 (2009).","journal-title":"Harmful Algae"},{"key":"3115_CR39","doi-asserted-by":"publisher","DOI":"10.1016\/j.hal.2023.102519","volume":"129","author":"SH Ahn","year":"2023","unstructured":"Ahn, S. H., Glibert, P. M. & Heil, C. A. In hot water: interactions of temperature, nitrogen form and availability and photosynthetic and nitrogen uptake responses in natural Karenia brevis populations. Harmful Algae 129, 102519 (2023).","journal-title":"Harmful Algae"},{"key":"3115_CR40","doi-asserted-by":"publisher","first-page":"615","DOI":"10.1038\/s41564-025-01948-2","volume":"10","author":"JR Seymour","year":"2025","unstructured":"Seymour, J. R. & McLellan, S. L. Climate change will amplify the impacts of harmful microorganisms in aquatic ecosystems. Nat. Microbiol. 10, 615\u2013626 (2025).","journal-title":"Nat. Microbiol."},{"key":"3115_CR41","doi-asserted-by":"publisher","DOI":"10.3389\/fevo.2021.646080","volume":"9","author":"CA Heil","year":"2021","unstructured":"Heil, C. A. & Muni-Morgan, A. L. Florida\u2019s harmful algal bloom (HAB) problem: escalating risks to human, environmental and economic health with climate change. Front. Ecol. Evol. 9, 646080 (2021).","journal-title":"Front. Ecol. Evol."},{"key":"3115_CR42","doi-asserted-by":"publisher","first-page":"280","DOI":"10.1038\/s41586-023-05760-y","volume":"615","author":"Y Dai","year":"2023","unstructured":"Dai, Y. et al. Coastal phytoplankton blooms expand and intensify in the 21st century. Nature 615, 280\u2013284 (2023).","journal-title":"Nature"},{"key":"3115_CR43","doi-asserted-by":"publisher","first-page":"844","DOI":"10.3389\/fmicb.2020.00844","volume":"11","author":"RG Hatfield","year":"2020","unstructured":"Hatfield, R. G. et al. The application of nanopore sequencing technology to the study of dinoflagellates: a proof of concept study for rapid sequence-based discrimination of potentially harmful algae. Front. Microbiol. 11, 844 (2020).","journal-title":"Front. Microbiol."},{"key":"3115_CR44","doi-asserted-by":"publisher","first-page":"555","DOI":"10.1080\/00288330.2017.1298632","volume":"51","author":"KF Smith","year":"2017","unstructured":"Smith, K. F., Kohli, G. S., Murray, S. A. & Rhodes, L. L. Assessment of the metabarcoding approach for community analysis of benthic\u2013epiphytic dinoflagellates using mock communities. NZ J. Mar. Freshw. Res. 51, 555\u2013576 (2017).","journal-title":"NZ J. Mar. Freshw. Res."},{"key":"3115_CR45","doi-asserted-by":"publisher","first-page":"1361","DOI":"10.3390\/md12031361","volume":"12","author":"KF Smith","year":"2014","unstructured":"Smith, K. F., De Salas, M., Adamson, J. & Rhodes, L. L. Rapid and accurate identification by real-time PCR of biotoxin-producing dinoflagellates from the family Gymnodiniaceae. Mar. Drugs 12, 1361\u20131376 (2014).","journal-title":"Mar. Drugs"},{"key":"3115_CR46","doi-asserted-by":"publisher","first-page":"5726","DOI":"10.1128\/AEM.69.9.5726-5730.2003","volume":"69","author":"M Gray","year":"2003","unstructured":"Gray, M., Wawrik, B., Paul, J. & Casper, E. Molecular detection and quantitation of the red tide dinoflagellate Karenia brevis in the marine environment. Appl. Environ. Microbiol. 69, 5726\u20135730 (2003).","journal-title":"Appl. Environ. Microbiol."},{"key":"3115_CR47","unstructured":"Yankovich, G. & Graham, N. Toxin caused by algae forces quarantine of Yorke Peninsula oysters. ABC News https:\/\/www.abc.net.au\/news\/2025-05-08\/millions-oysters-quarantined-sa-algal-bloom\/105266062 (2025)."},{"key":"3115_CR48","doi-asserted-by":"publisher","first-page":"563","DOI":"10.2216\/i0031-8884-42-6-563.1","volume":"42","author":"L Botes","year":"2003","unstructured":"Botes, L., Sym, S. D. & Pitcher, G. C. Karenia cristata sp. nov. and Karenia bicuneiformis sp. nov. (Gymnodiniales, Dinophyceae): two new Karenia species from the South African coast. Phycologia 42, 563\u2013571 (2003).","journal-title":"Phycologia"},{"key":"3115_CR49","doi-asserted-by":"publisher","first-page":"75","DOI":"10.1016\/j.hal.2014.10.006","volume":"40","author":"E N\u00e9zan","year":"2014","unstructured":"N\u00e9zan, E. et al. Genetic diversity of the harmful family Kareniaceae (Gymnodiniales, Dinophyceae) in France, with the description of Karlodinium gentienii sp. nov.: a new potentially toxic dinoflagellate. Harmful Algae 40, 75\u201391 (2014).","journal-title":"Harmful Algae"},{"key":"3115_CR50","doi-asserted-by":"publisher","first-page":"366","DOI":"10.1016\/j.hal.2011.01.002","volume":"10","author":"JJ Dorantes-Aranda","year":"2011","unstructured":"Dorantes-Aranda, J. J. et al. Novel application of a fish gill cell line assay to assess ichthyotoxicity of harmful marine microalgae. Harmful Algae 10, 366\u2013373 (2011).","journal-title":"Harmful Algae"},{"key":"3115_CR51","doi-asserted-by":"publisher","unstructured":"Rolton, A. et al. Comparative toxicity of Karlodinium veneficum from France and New Zealand on fish gill cells and oyster gametes. Harmful Algae 150, 102991 (2025); https:\/\/doi.org\/10.1016\/j.hal.2025.102991","DOI":"10.1016\/j.hal.2025.102991"},{"key":"3115_CR52","doi-asserted-by":"publisher","first-page":"560","DOI":"10.4319\/lo.2014.59.2.0560","volume":"59","author":"DR Hardison","year":"2014","unstructured":"Hardison, D. R., Sunda, W. G., Tester, P. A., Shea, D. & Litaker, W. R. Increased cellular brevetoxins in the red tide dinoflagellate Karenia brevis under CO2 limitation of growth rate: evolutionary implications and potential effects on bloom toxicity. Limnol. Oceanogr. 59, 560\u2013577 (2014).","journal-title":"Limnol. Oceanogr."},{"key":"3115_CR53","doi-asserted-by":"publisher","first-page":"621","DOI":"10.1289\/ehp.7499","volume":"113","author":"DG Baden","year":"2005","unstructured":"Baden, D. G., Bourdelais, A. J., Jacocks, H., Michelliza, S. & Naar, J. Natural and derivative brevetoxins: historical background, multiplicity, and effects. Environ. Health Perspect. 113, 621\u2013625 (2005).","journal-title":"Environ. Health Perspect."},{"key":"3115_CR54","doi-asserted-by":"publisher","first-page":"455","DOI":"10.1016\/j.toxicon.2004.02.017","volume":"43","author":"Z Wang","year":"2004","unstructured":"Wang, Z. et al. LC\/MS analysis of brevetoxin metabolites in the Eastern oyster (Crassostrea virginica). Toxicon 43, 455\u2013465 (2004).","journal-title":"Toxicon"},{"key":"3115_CR55","first-page":"611","volume":"47","author":"D Horstman","year":"1991","unstructured":"Horstman, D. et al. Red tides in False Bay, 1959\u20131989, with particular reference to recent blooms of Gymnodinium sp. Trans. R. Soc. S. Afr. 47, 611\u2013628 (1991).","journal-title":"Trans. R. Soc. S. Afr."},{"key":"3115_CR56","first-page":"1","volume":"15","author":"G Pitcher","year":"1996","unstructured":"Pitcher, G. & Matthews, S. Noxious Gymnodinium species in South African waters. Harmful Algae News 15, 1\u20133 (1996).","journal-title":"Harmful Algae News"},{"key":"3115_CR57","unstructured":"Lawrence, J. et al. Assessment and Management of Biotoxin Risks in Bivalve Molluscs. Technical Paper 551 (FAO Fisheries and Aquaculture, 2011)."},{"key":"3115_CR58","doi-asserted-by":"publisher","first-page":"82","DOI":"10.1016\/j.hal.2009.08.005","volume":"9","author":"B Kirkpatrick","year":"2010","unstructured":"Kirkpatrick, B. et al. Gastrointestinal emergency room admissions and Florida red tide blooms. Harmful Algae. 9, 82\u201386 (2010).","journal-title":"Harmful Algae."},{"key":"3115_CR59","unstructured":"CDC. Clinical signs and symptoms caused by saltwater harmful algal blooms. CDC https:\/\/www.cdc.gov\/harmful-algal-blooms\/hcp\/clinical-signs\/symptoms-saltwater-harmful-algal-blooms.html (2025)."},{"key":"3115_CR60","doi-asserted-by":"publisher","DOI":"10.1016\/j.hal.2020.101892","volume":"98","author":"JI Mardones","year":"2020","unstructured":"Mardones, J. I. et al. Unraveling the Karenia selliformis complex with the description of a non-gymnodimine producing Patagonian phylotype. Harmful Algae 98, 101892 (2020).","journal-title":"Harmful Algae"},{"key":"3115_CR61","doi-asserted-by":"publisher","first-page":"1026","DOI":"10.4319\/lo.2007.52.3.1026","volume":"52","author":"J Kubanek","year":"2007","unstructured":"Kubanek, J., Snell, T. W. & Pirkle, C. Chemical defense of the red tide dinoflagellate Karenia brevis against rotifer grazing. Limnol. Oceanogr. 52, 1026\u20131035 (2007).","journal-title":"Limnol. Oceanogr."},{"key":"3115_CR62","doi-asserted-by":"publisher","DOI":"10.1016\/j.aquatox.2020.105678","volume":"226","author":"X-D Li","year":"2020","unstructured":"Li, X.-D. et al. Progress on the usage of the rotifer Brachionus plicatilis in marine ecotoxicology: a review. Aquat. Toxicol. 226, 105678 (2020).","journal-title":"Aquat. Toxicol."},{"key":"3115_CR63","doi-asserted-by":"publisher","first-page":"149","DOI":"10.1080\/00288330.2011.616210","volume":"46","author":"F Shi","year":"2012","unstructured":"Shi, F. et al. The toxic effects of three dinoflagellate species from the genus Karenia on invertebrate larvae and finfish. NZ J. Mar. Freshw. Res. 46, 149\u2013165 (2012).","journal-title":"NZ J. Mar. Freshw. Res."},{"key":"3115_CR64","doi-asserted-by":"publisher","first-page":"166","DOI":"10.2216\/i0031-8884-43-2-166.1","volume":"43","author":"MF De Salas","year":"2004","unstructured":"De Salas, M. F., Bolch, C. J. & Hallegraeff, G. M. Karenia umbella sp. nov. (Gymnodiniales, Dinophyceae), a new potentially ichthyotoxic dinoflagellate species from Tasmania, Australia. Phycologia 43, 166\u2013175 (2004).","journal-title":"Phycologia"},{"key":"3115_CR65","doi-asserted-by":"publisher","DOI":"10.3389\/frpro.2024.1422481","volume":"2","author":"A Verma","year":"2024","unstructured":"Verma, A. et al. Characterization and inter-strain variability in ichthyotoxicity of Heterocapsa ovata (Peridiniales, Dinophyceae) from temperate waters of South Australia. Front. Protistol. 2, 1422481 (2024).","journal-title":"Front. Protistol."},{"key":"3115_CR66","unstructured":"Roberts, S. Fish Kill Investigation: Coffin Bay Harmful Algal (Karenia mikimotoi) Bloom February 2014 (Primary Industries and Regions South Australia, 2014)."},{"key":"3115_CR67","doi-asserted-by":"publisher","unstructured":"Schlegel, R. W. Marine Heatwave Tracker (Dalhousie Univ. & Ocean Frontier Institute, 2020); https:\/\/doi.org\/10.5281\/zenodo.3787872","DOI":"10.5281\/zenodo.3787872"},{"key":"3115_CR68","doi-asserted-by":"publisher","first-page":"4975","DOI":"10.1073\/pnas.1619575114","volume":"114","author":"CJ Gobler","year":"2017","unstructured":"Gobler, C. J. et al. Ocean warming since 1982 has expanded the niche of toxic algal blooms in the North Atlantic and North Pacific oceans. Proc. Natl Acad. Sci. USA 114, 4975\u20134980 (2017).","journal-title":"Proc. Natl Acad. Sci. USA"},{"key":"3115_CR69","doi-asserted-by":"publisher","DOI":"10.3389\/fmars.2022.939393","volume":"9","author":"S Takagi","year":"2022","unstructured":"Takagi, S. et al. Controlling factors of large-scale harmful algal blooms with Karenia selliformis after record-breaking marine heatwaves. Front. Mar. Sci. 9, 939393 (2022).","journal-title":"Front. Mar. Sci."},{"key":"3115_CR70","doi-asserted-by":"publisher","first-page":"695","DOI":"10.1038\/327695a0","volume":"327","author":"GW Lennon","year":"1987","unstructured":"Lennon, G. W. et al. Gravity currents and the release of salt from an inverse estuary. Nature 327, 695\u2013697 (1987).","journal-title":"Nature"},{"key":"3115_CR71","doi-asserted-by":"publisher","first-page":"1285","DOI":"10.1016\/S0278-4343(03)00129-8","volume":"23","author":"JR De Silva Samarasinghe","year":"2003","unstructured":"De Silva Samarasinghe, J. R., Bode, L. & Mason, L. B. Modelled response of Gulf St Vincent (South Australia) to evaporation, heating and winds. Cont. Shelf Res. 23, 1285\u20131313 (2003).","journal-title":"Cont. Shelf Res."},{"key":"3115_CR72","doi-asserted-by":"publisher","first-page":"335","DOI":"10.5004\/dwt.2009.264","volume":"2","author":"J K\u00e4mpf","year":"2009","unstructured":"K\u00e4mpf, J., Brokensha, C. & Bolton, T. Hindcasts of the fate of desalination brine in large inverse estuaries: Spencer Gulf and Gulf St. Vincent, South Australia. Desalin. Water Treat. 2, 335\u2013344 (2009).","journal-title":"Desalin. Water Treat."},{"key":"3115_CR73","doi-asserted-by":"publisher","first-page":"101702","DOI":"10.1016\/j.hal.2019.101702","volume":"90","author":"X Li","year":"2019","unstructured":"Li, X., Yan, T., Yu, R. & Zhou, M. A review of Karenia mikimotoi: bloom events, physiology, toxicity and toxic mechanism. Harmful Algae 90, 101702 (2019).","journal-title":"Harmful Algae"},{"key":"3115_CR74","doi-asserted-by":"publisher","DOI":"10.1038\/s41598-018-27845-9","volume":"8","author":"RX Poulin","year":"2018","unstructured":"Poulin, R. X. et al. Karenia brevis allelopathy compromises the lipidome, membrane integrity, and photosynthesis of competitors. Sci. Rep. 8, 9572 (2018).","journal-title":"Sci. Rep."},{"key":"3115_CR75","doi-asserted-by":"publisher","DOI":"10.1016\/j.marenvres.2025.107374","volume":"211","author":"Z Liu","year":"2025","unstructured":"Liu, Z. et al. Cell density-dependent mutual growth inhibition in harmful dinoflagellate Karenia selliformis. Mar. Environ. Res. 211, 107374 (2025).","journal-title":"Mar. Environ. Res."},{"key":"3115_CR76","doi-asserted-by":"publisher","first-page":"445","DOI":"10.1038\/s43247-025-02421-y","volume":"6","author":"E Silva","year":"2025","unstructured":"Silva, E. et al. Warming and freshening coastal waters impact harmful algal bloom frequency in high latitudes. Commun. Earth Environ. 6, 445 (2025).","journal-title":"Commun. Earth Environ."},{"key":"3115_CR77","unstructured":"QGIS Development Team. QGIS Geographic Information System (Open Source Geospatial Foundation Project, 2026); http:\/\/qgis.org"},{"key":"3115_CR78","doi-asserted-by":"publisher","first-page":"6924","DOI":"10.1021\/acs.est.3c10502","volume":"58","author":"R Ruvindy","year":"2024","unstructured":"Ruvindy, R. et al. An on-farm workflow for predictive management of paralytic shellfish toxin-producing harmful algal blooms for the aquaculture industry. Environ. Sci. Technol. 58, 6924\u20136933 (2024).","journal-title":"Environ. Sci. Technol."},{"key":"3115_CR79","doi-asserted-by":"publisher","first-page":"C01011","DOI":"10.1029\/2011JC007395","volume":"117","author":"C Hu","year":"2012","unstructured":"Hu, C., Lee, Z. & Franz, B. Chlorophyll a algorithms for oligotrophic oceans: a novel approach based on three-band reflectance difference. J. Geophys. Res. Oceans 117, C01011 (2012).","journal-title":"J. Geophys. Res. Oceans"},{"key":"3115_CR80","doi-asserted-by":"publisher","first-page":"835","DOI":"10.1086\/160554","volume":"263","author":"JD Scargle","year":"1982","unstructured":"Scargle, J. D. Studies in astronomical time series analysis. II. Statistical aspects of spectral analysis of unevenly spaced data. Astrophys. J. 263, 835\u2013853 (1982).","journal-title":"Astrophys. J."},{"key":"3115_CR81","doi-asserted-by":"publisher","first-page":"16","DOI":"10.3847\/1538-4365\/aab766","volume":"236","author":"JT Van der Plas","year":"2018","unstructured":"Van der Plas, J. T. Understanding the Lomb\u2013Scargle periodogram. Astrophys. J. Suppl. Ser. 236, 16 (2018).","journal-title":"Astrophys. J. Suppl. Ser."},{"key":"3115_CR82","doi-asserted-by":"publisher","first-page":"633","DOI":"10.1111\/j.1529-8817.1987.tb04217.x","volume":"23","author":"MD Keller","year":"1987","unstructured":"Keller, M. D., Selvin, R. C., Claus, W. & Guillard, R. R. Media for the culture of oceanic ultraphytoplankton. J. Phycol. 23, 633\u2013638 (1987).","journal-title":"J. Phycol."},{"key":"3115_CR83","doi-asserted-by":"publisher","first-page":"999","DOI":"10.1111\/j.0022-3646.1994.00999.x","volume":"30","author":"CA Scholin","year":"1994","unstructured":"Scholin, C. A., Herzog, M., Sogin, M. & Anderson, D. M. Identification of group- and strain-specific genetic markers for globally distributed Alexandrium (Dinophyceae). J. Phycol. 30, 999\u20131011 (1994).","journal-title":"J. Phycol."},{"key":"3115_CR84","doi-asserted-by":"publisher","first-page":"10","DOI":"10.14806\/ej.17.1.200","volume":"17","author":"M Martin","year":"2011","unstructured":"Martin, M. Cutadapt removes adapter sequences from high-throughput sequencing reads. EMBnet J. 17, 10\u201312 (2011).","journal-title":"EMBnet J."},{"key":"3115_CR85","doi-asserted-by":"publisher","first-page":"581","DOI":"10.1038\/nmeth.3869","volume":"13","author":"B Callahan","year":"2016","unstructured":"Callahan, B. et al. DADA2: high resolution sample inference from Illumina amplicon data. Nat. Methods 13, 581\u2013583 (2016).","journal-title":"Nat. Methods"},{"key":"3115_CR86","doi-asserted-by":"publisher","DOI":"10.1371\/journal.pone.0061217","volume":"8","author":"PJ McMurdie","year":"2013","unstructured":"McMurdie, P. J. & Holmes, S. phyloseq: An R package for reproducible interactive analysis and graphics of microbiome census data. PLoS ONE 8, e61217 (2013).","journal-title":"PLoS ONE"},{"key":"3115_CR87","doi-asserted-by":"publisher","first-page":"411","DOI":"10.1111\/j.1529-8817.2005.04168.x","volume":"41","author":"H Zhang","year":"2005","unstructured":"Zhang, H., Bhattacharya, D. & Lin, S. Phylogeny of dinoflagellates based on mitochondrial cytochrome b and nuclear small subunit rDNA sequence comparisons. J. Phycol. 41, 411\u2013420 (2005).","journal-title":"J. Phycol."},{"key":"3115_CR88","doi-asserted-by":"publisher","first-page":"772","DOI":"10.1093\/molbev\/mst010","volume":"30","author":"K Katoh","year":"2013","unstructured":"Katoh, K. & Standley, D. M. MAFFT multiple sequence alignment software version 7: improvements in performance and usability. Mol. Biol. Evol. 30, 772\u2013780 (2013).","journal-title":"Mol. Biol. Evol."},{"key":"3115_CR89","doi-asserted-by":"publisher","first-page":"1972","DOI":"10.1093\/bioinformatics\/btp348","volume":"25","author":"S Capella-Guti\u00e9rrez","year":"2009","unstructured":"Capella-Guti\u00e9rrez, S., Silla-Mart\u00ednez, J. M. & Gabald\u00f3n, T. trimAl: A tool for automated alignment trimming in large-scale phylogenetic analyses. Bioinformatics 25, 1972\u20131973 (2009).","journal-title":"Bioinformatics"},{"key":"3115_CR90","doi-asserted-by":"publisher","first-page":"1530","DOI":"10.1093\/molbev\/msaa015","volume":"37","author":"BQ Minh","year":"2020","unstructured":"Minh, B. Q. et al. IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Mol. Biol. Evol. 37, 1530\u20131534 (2020).","journal-title":"Mol. Biol. Evol."},{"key":"3115_CR91","doi-asserted-by":"publisher","first-page":"539","DOI":"10.1093\/sysbio\/sys029","volume":"61","author":"F Ronquist","year":"2012","unstructured":"Ronquist, F. et al. MrBayes 3.2: efficient Bayesian phylogenetic inference and model choice across a large model space. Syst. Biol. 61, 539\u2013542 (2012).","journal-title":"Syst. Biol."},{"key":"3115_CR92","doi-asserted-by":"publisher","unstructured":"Murray, S. A. et al. A catastrophic marine mortality event caused by a complex algal bloom including the brevetoxin producer, Karenia cristata. Zenodo https:\/\/doi.org\/10.5281\/zenodo.20227729 (2026).","DOI":"10.5281\/zenodo.20227729"}],"container-title":["Nature Ecology & Evolution"],"original-title":[],"language":"en","link":[{"URL":"https:\/\/www.nature.com\/articles\/s41559-026-03115-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-026-03115-0","content-type":"text\/html","content-version":"vor","intended-application":"text-mining"},{"URL":"https:\/\/www.nature.com\/articles\/s41559-026-03115-0.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2026,8,5]],"date-time":"2026-08-05T15:02:50Z","timestamp":1785942170000},"score":1,"resource":{"primary":{"URL":"https:\/\/www.nature.com\/articles\/s41559-026-03115-0"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2026,7,6]]},"references-count":92,"journal-issue":{"issue":"8","published-online":{"date-parts":[[2026,8]]}},"alternative-id":["3115"],"URL":"https:\/\/doi.org\/10.1038\/s41559-026-03115-0","relation":{},"ISSN":["2397-334X"],"issn-type":[{"value":"2397-334X","type":"electronic"}],"subject":[],"published":{"date-parts":[[2026,7,6]]},"assertion":[{"value":"23 October 2025","order":1,"name":"received","label":"Received","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"2 June 2026","order":2,"name":"accepted","label":"Accepted","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"6 July 2026","order":3,"name":"first_online","label":"First Online","group":{"name":"ArticleHistory","label":"Article History"}},{"value":"The authors declare no competing interests.","order":1,"name":"Ethics","label":"Competing interests","group":{"name":"EthicsHeading","label":"Ethics"}}]}}