Philippa Kaur

Verma, A. et al. The genetic basis of toxin biosynthesis in dinofagellates. Microorganisms 7, 222 (2019).Article 
CAS 

Google Scholar 
Hallegraeff, G. M. Ocean climate change, phytoplankton community responses, and harmful algal blooms: A formidable predictive challenge1. J. Phycol. 46, 220–235 (2010).Article 
CAS 

Google Scholar 
Hoppenrath, M., Murray, S., Chomérat, N., Horiguchi, T. Marine Benthic Dinoflagellates – Unveiling Their Worldwide Biodiversity (Kleine Senckenberg-reihe 54). E. Schweizerbart’sche Verlagbuchhandlung (2014).Luo, Z. et al. Cryptic diversity within the harmful dinoflagellate Akashiwo sanguinea in coastal Chinese waters is related to differentiated ecological niches. Harmful Algae 66, 88–96 (2017).Article 

Google Scholar 
Litaker, R. W. et al. Taxonomy of Gambierdiscus including four new species, Gambierdiscus caribaeus, Gambierdiscus carolinianus, Gambierdiscus carpenteri and Gambierdiscus ruetzleri (Gonyaulacales, Dinophyceae). Phycologia 48, 344–390 (2009).Article 

Google Scholar 
Hoppenrath, M. et al. Taxonomy and phylogeny of the benthic Prorocentrum species (Dinophyceae)—A proposal and review. Harmful Algae 27, 1–28 (2013).Article 

Google Scholar 
Wells, M. L. et al. Future HAB science: Directions and challenges in a changing climate. Harmful Algae 91, 101632 (2020).Article 

Google Scholar 
Rhodes, L. World-wide occurrence of the toxic dinoflagellate genus Ostreopsis Schmidt. Toxicon 57, 400–407 (2011).Article 
CAS 

Google Scholar 
Parsons, M. L. et al. Gambierdiscus and Ostreopsis: Reassessment of the state of knowledge of their taxonomy, geography, ecophysiology, and toxicology. Harmful Algae 14, 107–129 (2012).Article 
CAS 

Google Scholar 
Schmidt, J. Preliminary report of the botanical results of the Danish expedition to Siam (1899–1900). Part IV Peridiniales. Bot. Tidsskr. 24, 212–221 (1901).
Google Scholar 
Accoroni, S. et al. Ostreopsis fattorussoi sp. nov. (Dinophyceae), a new benthic toxic Ostreopsis species from the eastern Mediterranean Sea. J. Phycol. 52, 1064–1084 (2016).Article 
CAS 

Google Scholar 
Verma, A., Hoppenrath, M., Dorantes-Aranda, J. J., Harwood, D. T. & Murray, S. A. Molecular and phylogenetic characterization of Ostreopsis (Dinophyceae) and the description of a new species, Ostreopsis rhodesae sp. nov., from a subtropical Australian lagoon. Harmful Algae 60, 116–130 (2016).Article 
CAS 

Google Scholar 
Fukuyo, Y. Taxonomical study on benthic dinoflagellates collected in coral reefs. Nippon Suisan Gakk. 47, 967–978 (1981).Article 

Google Scholar 
Faust, M. A. Three new Ostreopsis species (Dinophyceae): O. marinus sp. nov., O. belizeanus sp. nov., and O. caribbeanus sp. nov.. Phycologia 38, 92–99 (1999).Article 

Google Scholar 
Faust, M. A. & Morton, S. L. Morphology and ecology of the marine dinoflagellate Ostreopsis labens sp. nov. (Dinophyceae). J. Phycol. 31, 456–463 (1995).Article 

Google Scholar 
Chomérat, N., Bilien, G., Couté, A. & Quod, J.-P. Reinvestigation of Ostreopsis mascarenensis Quod (Dinophyceae, Gonyaulacales) from Reunion Island (SW Indian Ocean): Molecular phylogeny and emended description. Phycologia 59, 140–153 (2020).Article 

Google Scholar 
Boisnoir, A., Bilien, G., Lemée, R. & Chomérat, N. First insights on the diversity of the genus Ostreopsis (Dinophyceae, Gonyaulacales) in Guadeloupe Island, with emphasis on the phylogenetic position of O. heptagona. Eur. J. Protistol. 83, 125875 (2022).Article 

Google Scholar 
Chomérat, N. et al. Ostreopsis lenticularis Y. Fukuyo (Dinophyceae, Gonyaulacales) from French Polynesia (South Pacific Ocean): A revisit of its morphology, molecular phylogeny and toxicity. Harmful Algae 84, 95–111 (2019).Article 

Google Scholar 
Nguyen-Ngoc, L. et al. Morphological and genetic analyses of Ostreopsis (Dinophyceae, Gonyaulacales, Ostreopsidaceae) species from Vietnamese waters with a re-description of the type species, O. siamensis 1. J. Phycol. 57, 1059–1083 (2021).Article 

Google Scholar 
Faust, M. A. Observation of sand-dwelling toxic dinoflagellates (Dinophyceae) from widely differing sites, including two new species. J. Phycol. 31, 996–1003 (1995).Article 

Google Scholar 
David, H., Laza-Martínez, A., Miguel, I. & Orive, E. Broad distribution of Coolia monotis and restricted distribution of Coolia cf. canariensis (Dinophyceae) on the Atlantic coast of the Iberian Peninsula. Phycologia 53, 342–352 (2014).Article 

Google Scholar 
Rhodes, L. L. et al. Toxic dinoflagellates (Dinophyceae) from Rarotonga Cook Islands. Toxicon 56, 751–758 (2010).Article 
CAS 

Google Scholar 
Meunier, A. Coolia monotis sp. nov. in Mémoires du Musée Royal d’Histoire Naturelle de Belgique. Microplankton Mer Flamande, Méme partie—Les Péridiniens 8, 68–69 (1919).
Google Scholar 
Rhodes, L. et al. Epiphytic dinoflagellates in sub-tropical New Zealand, in particular the genus Coolia Meunier. Harmful Algae 34, 36–41 (2014).Article 

Google Scholar 
Rhodes, L., Adamson, J., Suzuki, T., Briggs, L. & Garthwaite, I. Toxic marine epiphytic dinoflagellates, Ostreopsis siamensis and Coolia monotis (Dinophyceae), in New Zealand. N. Z. J. Mar. Freshw. Res. 34, 371–383 (2000).Article 

Google Scholar 
Fraga, S., Penna, A., Bianconi, I., Paz, B. & Zapata, M. Coolia canariensis sp. nov. (Dinophyceae), a new nontoxic epiuphytic benthic dinoflagellate from the Canary Islands 1. J. Phycol. 44, 1060–1070 (2008).Article 
CAS 

Google Scholar 
Lindemann, E. Abteilung Peridineae (Dinoflagellate). In Die Natürlichen Pflanzenfamilien nebst ihren Gattungen und wichtigeren Arten insbesondere den Nutzpflanzen, 3–104 (1928).Biecheler, B. Recherches sur les Péridiniens. Bulletin biologique de France et de Belgique Supplement 36, 1–149 (1952).
Google Scholar 
Balech, E. Étude des dinoflagellés du sable de Roscoff. Revue Algologique, Nouvelle Serie 2, 29–52 (1956).

Google Scholar 
Mohammad-Noor, N. et al. Autecology and phylogeny of Coolia tropicalis and Coolia malayensis (Dinophyceae), with emphasis on taxonomy of C. tropicalis based on light microscopy, scanning electron microscopy and LSU r DNA 1. J. Phycol. 49, 536–545 (2013).Article 

Google Scholar 
Leaw, C. P., Lim, P. T., Cheng, K. W., Ng, B. K. & Usup, G. Morphology and molecular characterization of a new species of thecate benthic dinoflagellate, Coolia malayensis sp. nov. (Dinophyceae) 1. J. Phycol. 46, 162–171 (2010).Article 
CAS 

Google Scholar 
Ten-Hage, L., Turquet, J., Quod, J. & Couté, A. Coolia areolata sp. nov. (Dinophyceae), a new sand-dwelling dinoflagellate from the southwestern Indian Ocean. Phycologia 39, 377–383 (2000).Article 

Google Scholar 
Karafas, S., York, R. & Tomas, C. Morphological and genetic analysis of the Coolia monotis species complex with the introduction of two new species, Coolia santacroce sp. nov. and Coolia palmyrensis sp. nov. (Dinophyceae). Harmful Algae 46, 18–33 (2015).Article 
CAS 

Google Scholar 
David, H., Laza-Martínez, A., Rodríguez, F., Fraga, S. & Orive, E. Coolia guanchica sp. nov.(Dinophyceae) a new epibenthic dinoflagellate from the Canary Islands (NE Atlantic Ocean). Eur. J. Phycol. 55, 76–88 (2020).Article 
CAS 

Google Scholar 
Sato, S. et al. Phylogeography of Ostreopsis along west Pacific coast, with special reference to a novel clade from Japan. PLoS One 6, e27983 (2011).Article 
ADS 
CAS 

Google Scholar 
Penna, A. et al. Characterization of Ostreopsis and Coolia (Dinophyceae) isolates in the western Mediterranean Sea based on morphology, toxicity and internal transcribed spacer 5.8 S rDNA sequences. J. Phycol. 41, 212–225 (2005).Article 
CAS 

Google Scholar 
Tawong, W. et al. Distribution and molecular phylogeny of the dinoflagellate genus Ostreopsis in Thailand. Harmful Algae 37, 160–171 (2014).Article 

Google Scholar 
Faimali, M. et al. Toxic effects of harmful benthic dinoflagellate Ostreopsis ovata on invertebrate and vertebrate marine organisms. Mar. Environ. Res. 76, 97–107 (2012).Article 
CAS 

Google Scholar 
Tubaro, A. et al. Case definitions for human poisonings postulated to palytoxins exposure. Toxicon 57, 478–495 (2011).Article 
CAS 

Google Scholar 
Ciminiello, P. et al. Investigation of the toxin profile of Greek mussels Mytilus galloprovincialis by liquid chromatography mass spectrometry. Toxicon 47, 174–181 (2006).Article 
CAS 

Google Scholar 
Giussani, V. et al. Active role of the mucilage in the toxicity mechanism of the harmful benthic dinoflagellate Ostreopsis cf. ovata. Harmful Algae 44, 46–53 (2015).Article 
CAS 

Google Scholar 
Usami, M. et al. Palytoxin analogs from the dinoflagellate Ostreopsis siamensis. J. Am. Chem. Soc. 117, 5389–5390 (1995).Article 
CAS 

Google Scholar 
Ukena, T. et al. Structure elucidation of ostreocin D, a palytoxin analog isolated from the dinoflagellate Ostreopsis siamensis. Biosci. Biotechnol. Biochem. 65, 2585–2588 (2001).Article 
CAS 

Google Scholar 
Amzil, Z. et al. Ovatoxin-a and palytoxin accumulation in seafood in relation to Ostreopsis cf. ovata blooms on the French Mediterranean coast. Mar. Drugs 10, 477–496 (2012).Article 
CAS 

Google Scholar 
Ciminiello, P. et al. Unique toxin profile of a Mediterranean Ostreopsis cf. ovata strain: HR LC-MS n characterization of ovatoxin-f, a new palytoxin congener. Chem. Res. Toxicol. 25, 1243–1252 (2012).Article 
CAS 

Google Scholar 
Laza-Martinez, A., Orive, E. & Miguel, I. Morphological and genetic characterization of benthic dinoflagellates of the genera Coolia, Ostreopsis and Prorocentrum from the south-eastern Bay of Biscay. Eur. J. Phycol. 46, 45–65 (2011).Article 

Google Scholar 
Holmes, M. J., Lewis, R. J., Jones, A. & Hoy, A. W. W. Cooliatoxin, the first toxin from Coolia monotis (Dinophyceae). Nat. Toxins 3, 355–362 (1995).Article 
CAS 

Google Scholar 
Rhodes, L. L. & Thomas, A. E. Coolia monotis (Dinophyceae): A toxic epiphytic microalgal species found in New Zealand (Note). N. Z. J. Mar. Freshw. Res. 31, 139–141 (1997).Article 
CAS 

Google Scholar 
Tibiriçá, C. EJd. A. et al. Diversity and toxicity of the genus Coolia Meunier in Brazil, and detection of 44-methyl Gambierone in Coolia tropicalis. Toxins 12, 327 (2020).Article 

Google Scholar 
Tillmann, U., Hoppenrath, M. & Gottschling, M. Reliable determination of Prorocentrum micans Ehrenb. (Prorocentrales, Dinophyceae) based on newly collected material from the type locality. Eur. J. Phycol 54, 417–431 (2019).Article 
CAS 

Google Scholar 
Chomérat, N. et al. Taxonomy and toxicity of a bloom-forming Ostreopsis species (Dinophyceae, Gonyaulacales) in Tahiti island (South Pacific Ocean): One step further towards resolving the identity of O. siamensis. Harmful Algae 98, 101888 (2020).Article 

Google Scholar 
Rhodes, L. L. et al. The dinoflagellate genera Gambierdiscus and Ostreopsis from subtropical Raoul Island and North Meyer Island, Kermadec Islands. N. Z. J. Mar. Freshw. Res. 51, 490–504 (2017).Article 
CAS 

Google Scholar 
Penna, A. et al. A phylogeographical study of the toxic benthic dinoflagellate genus Ostreopsis Schmidt. J. Biogeogr. 37, 830–841 (2010).Article 

Google Scholar 
Zhang, H. et al. Morphology and molecular phylogeny of Ostreopsis cf. ovata and O. lenticularis (Dinophyceae) from Hainan Island South China Sea. Phycol. Res. 66, 3–14 (2018).Article 
ADS 
CAS 

Google Scholar 
Carnicer, O., García-Altares, M., Andree, K. B., Diogène, J. & Fernández-Tejedor, M. First evidence of Ostreopsis cf. ovata in the eastern tropical Pacific Ocean Ecuadorian coast. Bot. Mar. 59, 267–274 (2016).
Google Scholar 
Nascimento, S. M. et al. Ostreopsis cf. ovata (Dinophyceae) molecular phylogeny, morphology, and detection of ovatoxins in strains and field samples from Brazil. Toxins 12, 70 (2020).Article 
CAS 

Google Scholar 
Caron, D. A. et al. Defining DNA-based operational taxonomic units for microbial-eukaryote ecology. Appl. Environ. Microbiol. 75, 5797–5808 (2009).Article 
ADS 
CAS 

Google Scholar 
McManus, G. B. & Katz, L. A. Molecular and morphological methods for identifying plankton: What makes a successful marriage?. J. Plankton Res. 31, 1119–1129 (2009).Article 
CAS 

Google Scholar 
De Vargas, C. et al. Eukaryotic plankton diversity in the sunlit ocean. Science 348, 1261605 (2015).Article 

Google Scholar 
del Campo, J. et al. Ecological and evolutionary significance of novel protist lineages. Eur. J. Protistol. 55, 4–11 (2016).Article 

Google Scholar 
Hallegraeff, G. Harmful algal blooms: A global overview. Man. Harmful Mar. Microalgae 33, 1–22 (2003).
Google Scholar 
Penna, A., Casabianca, S., Guerra, A. F., Vernesi, C. & Scardi, M. Analysis of phytoplankton assemblage structure in the Mediterranean Sea based on high-throughput sequencing of partial 18S rRNA sequences. Mar. Genom. 36, 49–55 (2017).Article 

Google Scholar 
Zarauz, L. & Irigoien, X. Effects of Lugol’s fixation on the size structure of natural nano–microplankton samples, analyzed by means of an automatic counting method. J. Plankton Res. 30, 1297–1303 (2008).Article 

Google Scholar 
De Luca, D., Piredda, R., Sarno, D. & Kooistra, W. H. Resolving cryptic species complexes in marine protists: phylogenetic haplotype networks meet global DNA metabarcoding datasets. ISME J. 15, 1931–1942 (2021).Article 

Google Scholar 
Wang, Z. et al. Phytoplankton community and HAB species in the South China Sea detected by morphological and metabarcoding approaches. Harmful Algae 118, 102297 (2022).Article 
CAS 

Google Scholar 
Le Bescot, N. et al. Global patterns of pelagic dinoflagellate diversity across protist size classes unveiled by metabarcoding. Environ. Microbiol. 18, 609–626 (2016).Article 

Google Scholar 
Hoppenrath, M. Dinoflagellate taxonomy—A review and proposal of a revised classification. Mar. Biodivers. 47, 381–403 (2017).Article 

Google Scholar 
Boenigk, J., Ereshefsky, M., Hoef-Emden, K., Mallet, J. & Bass, D. Concepts in protistology: Species definitions and boundaries. Eur. J. Protistol. 48, 96–102 (2012).Article 

Google Scholar 
David, H., Laza-Martínez, A., Miguel, I. & Orive, E. Ostreopsis cf. siamensis and Ostreopsis cf. ovata from the Atlantic Iberian Peninsula: Morphological and phylogenetic characterization. Harmful Algae 30, 44–55 (2013).Article 
CAS 

Google Scholar 
Aligizaki, K. & Nikolaidis, G. The presence of the potentially toxic genera Ostreopsis and Coolia (Dinophyceae) in the North Aegean Sea Greece. Harmful Algae 5, 717–730 (2006).Article 

Google Scholar 
Selina, M. S. & Orlova, T. Y. First occurrence of the genus Ostreopsis (Dinophyceae) in the Sea of Japan. Bot. Mar. 53, 243–249 (2010).Article 

Google Scholar 
Kang, N. S. et al. Morphology and molecular characterization of the epiphytic benthic dinoflagellate Ostreopsis cf. ovata in the temperate waters off Jeju Island Korea. Harmful Algae 27, 98–112 (2013).Article 
CAS 

Google Scholar 
Momigliano, P., Sparrow, L., Blair, D. & Heimann, K. The diversity of Coolia spp. (Dinophyceae Ostreopsidaceae) in the central Great Barrier Reef region. PloS One 8, e79278 (2013).Article 
ADS 
CAS 

Google Scholar 
Nguyen, L. N. Morphology and distribution of the three epiphytic dinoflagellate species Coolia monotis, C. tropicalis, and C. canariensis (Ostreopsidaceae, Gonyaulacales, Dinophyceae) from Vietnamese coastal waters. Ocean Sci. 49, 211–221 (2014).Article 

Google Scholar 
Verma, A. et al. Functional significance of phylogeographic structure in a toxic benthic marine microbial eukaryote over a latitudinal gradient along the East Australian Current. Ecol. Evol. 10, 6257–6273 (2020).Article 

Google Scholar 
Wayne Litaker, R. et al. Recognizing dinoflagellate species using ITS rDNA sequences 1. J. Phycol. 43, 344–355 (2007).Article 

Google Scholar 
Kremp, A. et al. Phylogenetic relationships, morphological variation, and toxin patterns in the Alexandrium ostenfeldii (D inophyceae) complex: Implications for species boundaries and identities. J. Phycol. 50, 81–100 (2014).Article 
CAS 

Google Scholar 
Nascimento, S. M., da Silva, R. A., Oliveira, F., Fraga, S. & Salgueiro, F. Morphology and molecular phylogeny of Coolia tropicalis, Coolia malayensis and a new lineage of the Coolia canariensis species complex (Dinophyceae) isolated from Brazil. Eur. J. Phycol. 54, 484–496 (2019).Article 
CAS 

Google Scholar 
Phua, Y. H., Roy, M. C., Lemer, S., Husnik, F. & Wakeman, K. C. Diversity and toxicity of Pacific strains of the benthic dinoflagellate Coolia (Dinophyceae), with a look at the Coolia canariensis species complex. Harmful Algae 109, 102120 (2021).Article 

Google Scholar 
Selwood, A. I. et al. A sensitive assay for palytoxins, ovatoxins and ostreocins using LC-MS/MS analysis of cleavage fragments from micro-scale oxidation. Toxicon 60, 810–820 (2012).Article 
CAS 

Google Scholar 
Ciminiello, P. et al. Isolation and structure elucidation of ovatoxin-a, the major toxin produced by Ostreopsis ovata. J. Am. Chem. Soc. 134, 1869–1875 (2012).Article 
CAS 

Google Scholar 
Dell’Aversano, C. et al. Ostreopsis cf. ovata from the Mediterranean area. Variability in toxinprofiles and structural elucidation of unknowns through LC-HRMSn. In Proc. of the 16th International Conference on Harmful Algae, 70–73 (2014).Terajima, T., Uchida, H., Abe, N. & Yasumoto, T. Structure elucidation of ostreocin-A and ostreocin-E1, novel palytoxin analogs produced by the dinoflagellate Ostreopsis siamensis, using LC/Q-TOF MS. Biosci. Biotechnol. Biochem. 83, 381–390 (2019).Article 
CAS 

Google Scholar 
Tartaglione, L. et al. Chemical, molecular, and eco-toxicological investigation of Ostreopsis sp. from Cyprus Island: Structural insights into four new ovatoxins by LC-HRMS/MS. Anal. Bioanal. Chem. 408, 915–932 (2016).Article 
CAS 

Google Scholar 
Murray, J. S. et al. The role of 44-methylgambierone in ciguatera fish poisoning: Acute toxicity, production by marine microalgae and its potential as a biomarker for Gambierdiscus spp. Harmful Algae 97, 101853 (2020).Article 
CAS 

Google Scholar 
Nakajima, I., Oshima, Y. & Yasumoto, T. Toxicity of benthic dinoflagellates found in coral reef. Toxicity of benthic dinoflagellates in Okinawa. Nippon Suisan Gakk. 47, 1029–1033 (1981).Article 

Google Scholar 
Boente-Juncal, A. et al. Structure elucidation and biological evaluation of maitotoxin-3, a homologue of gambierone, from Gambierdiscus belizeanus. Toxins 11, 79 (2019).Article 
CAS 

Google Scholar 
Stuart, J. et al. Geographical distribution, molecular and toxin diversity of the dinoflagellate species Gambierdiscus honu in the Pacific region. Harmful Algae 118, 102308 (2022).Article 
CAS 

Google Scholar 
Smith, K. F. et al. A new Gambierdiscus species (Dinophyceae) from Rarotonga, Cook Islands: Gambierdiscus cheloniae sp. nov. Harmful Algae 60, 45–56 (2016).Article 
CAS 

Google Scholar 
Guillard, R. R. L. Culture of Marine Invertebrates Animals 29–60 (Plenum Press, 1975).Book 

Google Scholar 
Chomérat, N., iti Gatti, C. M., Nézan, É. & Chinain, M. Studies on the benthic genus Sinophysis (Dinophysales, Dinophyceae) II. S. canaliculata from Rapa Island (French Polynesia). Phycologia 56, 193–203 (2017).Article 

Google Scholar 
Abràmoff, M. D., Magalhães, P. J. & Ram, S. J. Image processing with ImageJ. Biophotonics Int. 11, 36–42 (2004).
Google Scholar 
Verma, A. et al. Molecular phylogeny, morphology and toxigenicity of Ostreopsis cf. siamensis (Dinophyceae) from temperate south-east Australia. Phycol. Res. 64, 146–159 (2016).Article 
CAS 

Google Scholar 
Kearse, M. et al. Geneious basic: An integrated and extendable desktop software platform for the organization and analysis of sequence data. Bioinformatics 28, 1647–1649 (2012).Article 

Google Scholar 
Kumar, S., Stecher, G. & Tamura, K. MEGA7: Molecular evolutionary genetics analysis version 7.0 for bigger datasets. Mol. Biol. Evol. 33, 1870–1874 (2016).Article 
CAS 

Google Scholar 
Posada, D. & Crandall, K. A. MODELTEST: Testing the model of DNA substitution. Bioinformatics 14, 817–818 (1998).Article 
CAS 

Google Scholar 
Ronquist, F. & Huelsenbeck, J. P. MrBayes 3: Bayesian phylogenetic inference under mixed models. Bioinformatics 19, 1572–1574 (2003).Article 
CAS 

Google Scholar 
Murray, J. S. et al. Acute toxicity of gambierone and quantitative analysis of gambierones produced by cohabitating benthic dinoflagellates. Toxins 13, 333 (2021).Article 
CAS 

Google Scholar 
Murray, J. S., Boundy, M. J., Selwood, A. I. & Harwood, D. T. Development of an LC-MS/MS method to simultaneously monitor maitotoxins and selected ciguatoxins in algal cultures and P-CTX-1B in fish. Harmful Algae 80, 80–87 (2018).Article 
CAS 

Google Scholar  More

Animal collection and holding for this project was conducted under Marine Research Permit RE-19–28 issued by the Ministry of Natural Resources, Environment, and Tourism of the Republic of Palau (10.03.2019), Marine Research/Collection Permit and Agreement 62 issued by the Koror State Government (08.10.2019), Queensland Government GBRMPA Marine Parks Permit G14/36689.1, Queensland Government DNPRSR Marine Parks Permits QS2014/MAN247 and QS2014/MAN247a, Queensland Government General Fisheries Permit 168991, Queensland Government DAFF Animal Ethics approval CA2013/11/733, approval by The Bahamas Department of Marine Resources, approval by the Animal Care Officer of both the University of Bremen and the Leibniz Centre for Tropical Marine Research (ZMT), and in accordance with UK and Germany animal care guidelines.Sample collectionWe collected fish carbonate samples at four study locations across three tropical and subtropical regions: Eleuthera (24°50’N, 76°20’W), The Bahamas, between 2009 and 201127,37; Heron Reef (23°27’S, 151°55’E) and Moreton Bay (27°29’S, 153°24’E) in Queensland, Australia, in 2014 and 201528; and Koror (7°20’N, 134°28’E), Palau, during November and December 2019. These are located within four distinct marine biogeographic provinces and three realms (Tropical Atlantic, Central Indo-Pacific, and Temperate Australasia)43. At each location fish were collected using barrier nets, dip nets, clove oil or hook and line, and immediately transferred to aquaria facilities at the Cape Eleuthera Institute, Heron Island and Moreton Bay Research Stations, and the Palau International Coral Reef Center. Fish were held in a range of tanks (60, 400, or 1400 L in the Bahamas, 10, 60, 100, 120, or 400 L in Heron Island and Moreton Bay, and 8, 80, 280, or 400 L in Palau) of suitable dimensions for different fish sizes ( 5). Each sample was titrated with 0.01–0.5 N HCl (with continuous aeration with CO2-free air) until the end point (grey-lavender; pH~4.80) was reached and stable for at least 10 min. If the sample was over-titrated (pink), 0.01–0.1 N NaOH was added to titrate back to the end point and the amount of base used was subtracted from the amount of acid. Acid and base were added using an electronic multi-dispenser pipette (Eppendorf Repeater ®E3X, Eppendorf, Hamburg, Germany) with a precision of  ± 1 ({{{{{rm{mu }}}}}})L. Additionally, the pH of several samples was monitored using a pH microelectrode (Mettler Toledo InLab Micro) to ascertain the correctness of the colorimetric end point. The amount of carbonate in the sample was then calculated using Eq. (1). The method was validated using certified reference material (Alkalinity Standard Solution, 25,000 mg/L as CaCO3, HACH) and the accuracy in the determination of solid samples was verified using certified CaCO3 powder (Suprapur, ≥ 99.95% purity, Merck) samples (60–500 ({{{{{rm{mu }}}}}})g) and resulted in 96.53 ± 1.94% accuracy (mean ± SE; n = 8).To compare values obtained with the two titration methods we further analysed 12 samples collected at Lizard Island, Australia, in February 2016. Samples were collected at 24 h intervals from one individual of Lethrinus atkinsoni (f. Lethrinidae, body mass: 245 g), a group of five Lutjanus fulvus (f. Lutjanidae, mean body mass: 21 g), and an individual of Cephalopholis cyanostigma (f. Serranidae, body mass: 295 g), following the procedures described above. During sample collection water temperature ranged from 29.1 °C during the night to 32.6 °C during the day, with an average of ~31 °C, mean salinity was 35.4, and pHNBS ranged from 8.13 to 8.21. To compare the amount of carbonate measured by the two methods we added carbonate samples to 20 ml ultrapure water and disaggregated crystals via sonication. We then used a Metrohm Titrando autotitrator and Metrohm Aquatrode pH electrode to measure initial pH of the suspension of carbonates, then titrated each sample of carbonate in two stages. Firstly, they were titrated down to pH 4.80 using 0.1 M HCl, adding 20 µl increments of acid until this was sufficient to keep pH below 4.80 for 10 min whilst bubbling with CO2-free air. This first stage was comparable to the single end point titration used for samples collected in Palau. Secondly, whilst continuing to bubble with CO2-free air, further acid was added to the sample until it reached pH 3.89 and was stable for 1 min. Then 0.1 M NaOH was added to the samples to return them to the initial pH. For all samples the first end point titration (to pH 4.80) yielded slightly higher values for carbonate content than the second double titration. The ratio between the two methods (single end point/double titration) was 1.08 ± 0.01 (mean ± SE; range: 1.04–1.14; Supplementary Table 2). As we found a small but consistent difference between the two methods, all following analyses were initially performed on the actual data obtained with the double titration for samples from Australia and The Bahamas, and the single end point titration for samples from Palau. Then, to assess the robustness of the results, we repeated the analyses after applying a correction factor of 1.08 to the excretion rates of Palauan fishes (that used the single end point titration method). All results were consistent and robust to the measured difference between the titration methods (Supplementary Figs. 8, 9).Finally, measurements of multiple samples from each individual collected over periods of 18–169 h (median: 64 h) were combined to produce an average individual excretion rate in ({{{{{rm{mu }}}}}})mol h−1. For fish held in groups, carbonate excretion rates per individual (of average biomass) were obtained by averaging the total excretion rate of the group across the sampling period and dividing it by the number of individuals in the tank. Excretion rates obtained from fish groups thus evened the intraspecific variability within tanks, and are therefore more robust than those directly obtained from fish held individually. This aspect was considered in our models by fitting weighted regressions (see the “Statistical modelling” section). In total, we measured the carbonate excretion rates of 382 individual fishes arranged in 192 groups (i.e., independent observations), representing 85 species from 35 families across three tropical regions (180 individuals from 29 species in Australia, 90 individuals from 10 species in the Bahamas, and 112 individuals from 46 species in Palau; Supplementary Table 1).We assume that during the sampling of carbonates fishes were close to their resting metabolic rate and that their carbonate excretion rates are representative of fish at rest. Although the ratio of tank volume to fish volume in our study (median ~660; inter-quartile range ~180–1700) typically greatly exceeds the guideline ideal range for measuring resting metabolic rate (20–50)85, fishes were fasted prior to and throughout sampling, and in most instances their movement was somewhat constrained by tank volume. Fasting reduces metabolic rate in all animals, including fish, as they do not undergo energy-intensive digestive processes and use energy reserves to support vital processes, triggering metabolic changes in many tissues and reducing activity levels86,87. Additionally, other than the carbonate syphoning ( More

Benca, J. P., Duijnstee, I. A. & Looy, C. V. Fossilized pollen malformations as indicators of past environmental stress and meiotic disruption: Insights from modern conifers. Paleobiology, 1–34 (2022).Marshall, J. E., Lakin, J., Troth, I. & Wallace-Johnson, S. M. Uv-b radiation was the devonian-carboniferous boundary terrestrial extinction kill mechanism. Sci. Adv. 6, eaba0768 (2020).Article 
ADS 
CAS 

Google Scholar 
Looy, C. V., Twitchett, R. J., Dilcher, D. L., Van Konijnenburg-Van Cittert, J. H. & Visscher, H. Life in the end-permian dead zone. Proc. Natl. Acad. Sci. 98, 7879–7883 (2001).Article 
ADS 
CAS 

Google Scholar 
Foster, C. & Afonin, S. Abnormal pollen grains: An outcome of deteriorating atmospheric conditions around the permian-triassic boundary. J. Geol. Soc. 162, 653–659 (2005).Article 
ADS 

Google Scholar 
Hochuli, P. A., Schneebeli-Hermann, E., Mangerud, G. & Bucher, H. Evidence for atmospheric pollution across the permian-triassic transition. Geology 45, 1123–1126 (2017).Article 
ADS 

Google Scholar 
Galasso, F., Bucher, H. & Schneebeli-Hermann, E. Mapping monstrosity: Malformed sporomorphs across the smithian/spathian boundary interval and beyond (salt range, pakistan). Global Planet. Change 219, 103975 (2022).Article 

Google Scholar 
Van de Schootbrugge, B. et al. Floral changes across the triassic/jurassic boundary linked to flood basalt volcanism. Nat. Geosci. 2, 589–594 (2009).Article 
ADS 

Google Scholar 
Lindström, S. et al. Volcanic mercury and mutagenesis in land plants during the end-triassic mass extinction. Sci. Adv. 5, eaaw4018 (2019).Article 
ADS 

Google Scholar 
Gravendyck, J., Schobben, M., Bachelier, J. B. & Kürschner, W. M. Macroecological patterns of the terrestrial vegetation history during the end-triassic biotic crisis in the central european basin: A palynological study of the bonenburg section (nw-germany) and its supra-regional implications. Global Planet. Change 194, 103286 (2020).Article 

Google Scholar 
Vilas-Boas, M., Pereira, Z., Cirilli, S., Duarte, L. V. & Fernandes, P. New data on the palynology of the triassic-jurassic boundary of the silves group, lusitanian basin, portugal. Rev. Palaeobot. Palynol. 290, 104426 (2021).Article 

Google Scholar 
Galasso, F., Feist-Burkhardt, S. & Schneebeli-Hermann, E. The palynology of the toarcian oceanic anoxic event at dormettingen, southwest germany, with emphasis on changes in vegetational dynamics. Rev. Palaeobotany Palynol. 304, 104701 (2022).Article 

Google Scholar 
Galasso, F., Feist-Burkhardt, S. & Schneebeli-Hermann, E. Do spores herald the toarcian oceanic anoxic event?. Rev. Palaeobot. Palynol. 306, 104748 (2022).Article 

Google Scholar 
Hay, W. W. & Floegel, S. New thoughts about the cretaceous climate and oceans. Earth Sci. Rev. 115, 262–272 (2012).Article 
ADS 
CAS 

Google Scholar 
Faucher, G., Erba, E., Bottini, C. & Gambacorta, G. Calcareous nannoplankton response to the latest cenomanian oceanic anoxic event 2 perturbation. RIVISTA ITALIANA DI PALEONTOLOGIA E STRATIGRAFIA (2017).Cohen, K. M., Finney, S. C., Gibbard, P. L. & Fan, J.-X. The ics international chronostratigraphic chart. Epis. J. Int. Geosci. 36, 199–204 (2013).
Google Scholar 
Caron, M. & Homewood, P. Evolution of early planktic foraminifers. Mar. Micropaleontol. 7, 453–462 (1983).Article 
ADS 

Google Scholar 
Jarvis, I. et al. Microfossil assemblages and the cenomanian-turonian (late cretaceous) oceanic anoxic event. Cretac. Res. 9, 3–103 (1988).Article 

Google Scholar 
Huber, B. T., Leckie, R. M., Norris, R. D., Bralower, T. J. & CoBabe, E. Foraminiferal assemblage and stable isotopic change across the cenomanian-turonian boundary in the subtropical north atlantic. J. Foraminiferal Res. 29, 392–417 (1999).
Google Scholar 
Culver, S. J. & Rawson, P. F. Biotic response to global change: The last 145 million years (Cambridge University Press, 2006).Erba, E. Calcareous nannofossils and mesozoic oceanic anoxic events. Mar. Micropaleontol. 52, 85–106 (2004).Article 
ADS 

Google Scholar 
Gebhardt, H., Kuhnt, W. & Holbourn, A. Foraminiferal response to sea level change, organic flux and oxygen deficiency in the cenomanian of the tarfaya basin, southern morocco. Mar. Micropaleontol. 53, 133–157 (2004).Article 
ADS 

Google Scholar 
Hardenbol, J. et al. Mesozoic and cenozoic sequence chronostratigraphic framework of european basins. Soc. Sediment. Geol. (1998).Miller, K. G. et al. The phanerozoic record of global sea-level change. Science 310, 1293–1298 (2005).Article 
ADS 
CAS 

Google Scholar 
Voigt, S., Gale, A. S. & Voigt, T. Sea-level change, carbon cycling and palaeoclimate during the late cenomanian of northwest europe; an integrated palaeoenvironmental analysis. Cretac. Res. 27, 836–858 (2006).Article 

Google Scholar 
Haq, B. U. Cretaceous eustasy revisited. Global Planet. Change 113, 44–58 (2014).Article 
ADS 

Google Scholar 
Sames, B. et al. Short-term sea-level changes in a greenhouse world-a view from the cretaceous. Palaeogeogr. Palaeoclimatol. Palaeoecol. 441, 393–411 (2016).Article 

Google Scholar 
Arthur, M. A., Dean, W. E. & Pratt, L. M. Geochemical and climatic effects of increased marine organic carbon burial at the cenomanian/turonian boundary. Nature 335, 714–717 (1988).Article 
ADS 

Google Scholar 
Tsikos, H. et al. Carbon-isotope stratigraphy recorded by the cenomanian-turonian oceanic anoxic event: Correlation and implications based on three key localities. J. Geol. Soc. 161, 711–719 (2004).Article 
ADS 
CAS 

Google Scholar 
Jarvis, I., Lignum, J. S., Gröcke, D. R., Jenkyns, H. C. & Pearce, M. A. Black shale deposition, atmospheric co2 drawdown, and cooling during the cenomanian-turonian oceanic anoxic event. Paleoceanography26 (2011).van Bentum, E. C., Reichart, G.-J. & Damsté, J. S. S. Organic matter provenance, palaeoproductivity and bottom water anoxia during the cenomanian/turonian oceanic anoxic event in the newfoundland basin (northern proto north atlantic ocean). Org. Geochem. 50, 11–18 (2012).Article 

Google Scholar 
Owens, J. D., Lyons, T. W. & Lowery, C. M. Quantifying the missing sink for global organic carbon burial during a cretaceous oceanic anoxic event. Earth Planet. Sci. Lett. 499, 83–94 (2018).Article 
ADS 
CAS 

Google Scholar 
Bralower, T. J. Calcareous nannofossil biostratigraphy and assemblages of the cenomanian-turonian boundary interval: Implications for the origin and timing of oceanic anoxia. Paleoceanography 3, 275–316 (1988).Article 
ADS 

Google Scholar 
Leckie, R. M., Bralower, T. J. & Cashman, R. Oceanic anoxic events and plankton evolution: Biotic response to tectonic forcing during the mid-cretaceous. Paleoceanography 17, 13–1 (2002).Article 

Google Scholar 
Slater, S. M., Bown, P., Twitchett, R. J., Danise, S. & Vajda, V. Global record of “ghost’’ nannofossils reveals plankton resilience to high co2 and warming. Science 376, 853–856 (2022).Article 
ADS 
CAS 

Google Scholar 
Forster, A., Schouten, S., Moriya, K., Wilson, P. A. & Sinninghe Damsté, J. S. Tropical warming and intermittent cooling during the cenomanian/turonian oceanic anoxic event 2: Sea surface temperature records from the equatorial atlantic. Paleoceanography22 (2007).Barclay, R. S., McElwain, J. C. & Sageman, B. B. Carbon sequestration activated by a volcanic co2 pulse during ocean anoxic event 2. Nat. Geosci. 3, 205–208 (2010).Article 
ADS 
CAS 

Google Scholar 
Damsté, J. S. S., van Bentum, E. C., Reichart, G.-J., Pross, J. & Schouten, S. A co2 decrease-driven cooling and increased latitudinal temperature gradient during the mid-cretaceous oceanic anoxic event 2. Earth Planet. Sci. Lett. 293, 97–103 (2010).Article 
ADS 

Google Scholar 
Heimhofer, U. et al. Vegetation response to exceptional global warmth during oceanic anoxic event 2. Nat. Commun. 9, 1–8 (2018).Article 
CAS 

Google Scholar 
Huber, B. T., MacLeod, K. G., Watkins, D. K. & Coffin, M. F. The rise and fall of the cretaceous hot greenhouse climate. Global Planet. Change 167, 1–23 (2018).Article 
ADS 

Google Scholar 
Robinson, S. A. et al. Southern hemisphere sea-surface temperatures during the cenomanian-turonian: Implications for the termination of oceanic anoxic event 2. Geology 47, 131–134 (2019).Article 
ADS 
CAS 

Google Scholar 
Voigt, S., Gale, A. S. & Flögel, S. Midlatitude shelf seas in the cenomanian-turonian greenhouse world: Temperature evolution and north atlantic circulation. Paleoceanography19 (2004).Van Helmond, N. et al. Freshwater discharge controlled deposition of cenomanian-turonian black shales on the nw european epicontinental shelf (wunstorf, north germany). Clim. Past Discuss 10, 3755–3786 (2014).ADS 

Google Scholar 
Li, Y.-X., Montanez, I. P., Liu, Z. & Ma, L. Astronomical constraints on global carbon-cycle perturbation during oceanic anoxic event 2 (oae2). Earth Planet. Sci. Lett. 462, 35–46 (2017).Article 
ADS 
CAS 

Google Scholar 
O’Brien, C. L. et al. Cretaceous sea-surface temperature evolution: Constraints from tex86 and planktonic foraminiferal oxygen isotopes. Earth Sci. Rev. 172, 224–247 (2017).Article 
ADS 

Google Scholar 
Jones, C. E. & Jenkyns, H. C. Seawater strontium isotopes, oceanic anoxic events, and seafloor hydrothermal activity in the jurassic and cretaceous. Am. J. Sci. 301, 112–149 (2001).Article 
ADS 
CAS 

Google Scholar 
Snow, L. J., Duncan, R. A. & Bralower, T. J. Trace element abundances in the rock canyon anticline, pueblo, colorado, marine sedimentary section and their relationship to caribbean plateau construction and oxygen anoxic event 2. Paleoceanography20 (2005).Kuroda, J. et al. Contemporaneous massive subaerial volcanism and late cretaceous oceanic anoxic event 2. Earth Planet. Sci. Lett. 256, 211–223 (2007).Article 
ADS 
CAS 

Google Scholar 
Turgeon, S. C. & Creaser, R. A. Cretaceous oceanic anoxic event 2 triggered by a massive magmatic episode. Nature 454, 323–326 (2008).Article 
ADS 
CAS 

Google Scholar 
Floegel, S. et al. Simulating the biogeochemical effects of volcanic co2 degassing on the oxygen-state of the deep ocean during the cenomanian/turonian anoxic event (oae2). Earth Planet. Sci. Lett. 305, 371–384 (2011).Article 
ADS 
CAS 

Google Scholar 
Tegner, C. et al. Magmatism and eurekan deformation in the high arctic large igneous province: 40ar-39ar age of kap washington group volcanics, north greenland. Earth Planet. Sci. Lett. 303, 203–214 (2011).Article 
ADS 
CAS 

Google Scholar 
Du Vivier, A. D. et al. Marine 187os/188os isotope stratigraphy reveals the interaction of volcanism and ocean circulation during oceanic anoxic event 2. Earth Planet. Sci. Lett. 389, 23–33 (2014).Article 
ADS 

Google Scholar 
Du Vivier, A., Selby, D., Condon, D., Takashima, R. & Nishi, H. Pacific 187os/188os isotope chemistry and u-pb geochronology: Synchroneity of global os isotope change across oae 2. Earth Planet. Sci. Lett. 428, 204–216 (2015).Article 
ADS 

Google Scholar 
Meyers, P. A. Why are the (delta )13corg values in phanerozoic black shales more negative than in modern marine organic matter?. Geochem. Geophys. Geosyst. 15, 3085–3106 (2014).Article 
ADS 
CAS 

Google Scholar 
Jenkyns, H. C., Dickson, A. J., Ruhl, M. & Van den Boorn, S. H. Basalt-seawater interaction, the plenus cold event, enhanced weathering and geochemical change: Deconstructing oceanic anoxic event 2 (cenomanian-turonian, late cretaceous). Sedimentology 64, 16–43 (2017).Article 
CAS 

Google Scholar 
Scaife, J. et al. Sedimentary mercury enrichments as a marker for submarine large igneous province volcanism? evidence from the mid-cenomanian event and oceanic anoxic event 2 (late cretaceous). Geochem. Geophys. Geosyst. 18, 4253–4275 (2017).Article 
ADS 
CAS 

Google Scholar 
Schröder-Adams, C. J., Herrle, J. O., Selby, D., Quesnel, A. & Froude, G. Influence of the high arctic igneous province on the cenomanian/turonian boundary interval, sverdrup basin, high canadian arctic. Earth Planet. Sci. Lett. 511, 76–88 (2019).Article 
ADS 

Google Scholar 
Jolet, P., Philip, J., Thomel, G., Lopez, G. & Tronchetti, G. Nouvelles données biostratigraphiques sur la limite cénomanien-turonien. la coupe de cassis (sud-est de la france): Proposition d’un hypostratotype européen. Comptes Rendus de l’Académie des Sciences-Series IIA-Earth and Planetary Science325, 703–709 (1997).Bown, P. R. & Young, J. Calcareous nannofossil biostratigraphy (Springer, 1998).Green, T., Renne, P. R. & Keller, C. B. Continental flood basalts drive phanerozoic extinctions. Proc. Natl. Acad. Sci. 119, e2120441119 (2022).Article 
CAS 

Google Scholar 
Percival, L. M. et al. Does large igneous province volcanism always perturb the mercury cycle? Comparing the records of oceanic anoxic event 2 and the end-cretaceous to other mesozoic events. Am. J. Sci. 318, 799–860 (2018).Article 
ADS 
CAS 

Google Scholar 
Salazar, L. et al. Diversity patterns of ferns along elevational gradients in andean tropical forests. Plant Ecol. Divers. 8, 13–24 (2015).Article 

Google Scholar 
Mehltreter, K., Walker, L. R. & Sharpe, J. M. Fern ecology (Cambridge University Press, 2010).Carvajal-Hernández, C. I., Gómez-Díaz, J. A., Kessler, M. & Krömer, T. Influence of elevation and habitat disturbance on the functional diversity of ferns and lycophytes. Plant Ecol. Divers. 11, 335–347 (2018).Article 

Google Scholar 
Kürschner, W. M., Batenburg, S. J. & Mander, L. Aberrant classopollis pollen reveals evidence for unreduced (2 n) pollen in the conifer family cheirolepidiaceae during the triassic-jurassic transition. Proc. Royal Soc. B: Biol. Sci. 280, 20131708 (2013).Article 

Google Scholar 
Traverse, A. Paleopalynology Vol. 28 (Springer Science & Business Media, 2007).Tyson, R. V. Palynofacies investigation of callovian (middle jurassic) sediments from dsdp site 534, blake-bahama basin, western central atlantic. Mar. Pet. Geol. 1, 3–13 (1984).Article 

Google Scholar 
RV, T. Sedimentary organic matter: Organic facies and palynofacieschapman & hall. London, 615pp (1995).Vakhrameyev, V. Classopollis pollen as an indicator of jurassic and cretaceous climate. Int. Geol. Rev. 24, 1190–1196 (1982).Article 

Google Scholar 
Vakhrameev, V. Range and palaeoecology of mesozoic conifers, the cheirolepidiaceae. Paleontol. Zh. 1, 19–34 (1970).
Google Scholar 
WATSON, J. Some lower cretaceous conifers of the cheirolepidiaceae from the usa and england. Palaeontology 20, 715–749 (1977).
Google Scholar 
Fonseca, C., Mendonça Filho, J. G., Lézin, C., De Oliveira, A. D. & Duarte, L. V. Organic matter deposition and paleoenvironmental implications across the cenomanian-turonian boundary of the subalpine basin (se france): Local and global controls. Int. J. Coal Geol. 218, 103364 (2020).Article 
CAS 

Google Scholar 
Benca, J. P., Duijnstee, I. A. & Looy, C. V. Uv-b-induced forest sterility: Implications of ozone shield failure in earth’s largest extinction. Sci. Adv. 4, e1700618 (2018).Article 
ADS 

Google Scholar 
Wilson, L. A study in variation of picea glauca (moench) voss pollen. Grana 4, 380–387 (1963).
Google Scholar 
Lindström, S., McLoughlin, S. & Drinnan, A. N. Intraspecific variation of taeniate bisaccate pollen within permian glossopterid sporangia, from the prince charles mountains, antarctica. Int. J. Plant Sci. 158, 673–684 (1997).Article 

Google Scholar 
Leitch, A. & Leitch, I. Ecological and genetic factors linked to contrasting genome dynamics in seed plants. New Phytol. 194, 629–646 (2012).Article 
CAS 

Google Scholar 
Coffin, M. F. & Eldholm, O. Large igneous provinces: crustal structure, dimensions, and external consequences. Rev. Geophys. 32, 1–36 (1994).Article 
ADS 

Google Scholar 
Wignall, P. B. Large igneous provinces and mass extinctions. Earth Sci. Rev. 53, 1–33 (2001).Article 
ADS 
CAS 

Google Scholar 
McElwain, J. C., Wade-Murphy, J. & Hesselbo, S. P. Changes in carbon dioxide during an oceanic anoxic event linked to intrusion into gondwana coals. Nature 435, 479–482 (2005).Article 
ADS 
CAS 

Google Scholar 
Bond, D. P., Wignall, P. B., Keller, G. & Kerr, A. Large igneous provinces and mass extinctions: An update. Volcan., Impacts, Mass Extinc.: Causes Effects 505, 29–55 (2014).
Google Scholar 
Burgess, S., Bowring, S., Fleming, T. & Elliot, D. High-precision geochronology links the ferrar large igneous province with early-jurassic ocean anoxia and biotic crisis. Earth Planet. Sci. Lett. 415, 90–99 (2015).Article 
ADS 
CAS 

Google Scholar 
Burgess, S. D., Muirhead, J. D. & Bowring, S. A. Initial pulse of siberian traps sills as the trigger of the end-permian mass extinction. Nat. Commun. 8, 1–6 (2017).Article 
ADS 
CAS 

Google Scholar 
Ernst, R. E. & Youbi, N. How large igneous provinces affect global climate, sometimes cause mass extinctions, and represent natural markers in the geological record. Palaeogeogr. Palaeoclimatol. Palaeoecol. 478, 30–52 (2017).Article 

Google Scholar 
Ruhl, M. et al. Reduced plate motion controlled timing of early jurassic karoo-ferrar large igneous province volcanism. Sci. Adv. 8, eabo0866 (2022).Article 
CAS 

Google Scholar 
Dickens, G. R., Paull, C. K. & Wallace, P. Direct measurement of in situ methane quantities in a large gas-hydrate reservoir. Nature 385, 426–428 (1997).Article 
ADS 
CAS 

Google Scholar 
Courtillot, V. E. & Renne, P. R. On the ages of flood basalt events. C.R. Geosci. 335, 113–140 (2003).Article 
ADS 

Google Scholar 
Rampino, M. R., Rodriguez, S., Baransky, E. & Cai, Y. Global nickel anomaly links siberian traps eruptions and the latest permian mass extinction. Sci. Rep. 7, 1–6 (2017).Article 
CAS 

Google Scholar 
Clapham, M. E. & Renne, P. R. Flood basalts and mass extinctions. Annu. Rev. Earth Planet. Sci. 47, 275–303 (2019).Article 
ADS 
CAS 

Google Scholar 
McElwain, J. C., Popa, M. E., Hesselbo, S. P., Haworth, M. & Surlyk, F. Macroecological responses of terrestrial vegetation to climatic and atmospheric change across the triassic/jurassic boundary in east greenland. Paleobiology 33, 547–573 (2007).Article 

Google Scholar 
Van de Schootbrugge, B. et al. End-triassic calcification crisis and blooms of organic-walled ‘disaster species’. Palaeogeogr. Palaeoclimatol. Palaeoecol. 244, 126–141 (2007).Article 

Google Scholar 
Ruckwied, K., Götz, A. E., Pálfy, J. & Török, Á. Palynology of a terrestrial coal-bearing series across the triassic/jurassic boundary (mecsek mts, hungary). Central Euro. Geol. 51, 1–15 (2008).Article 
CAS 

Google Scholar 
Götz, A., Ruckwied, K., Pálfy, J. & Haas, J. Palynological evidence of synchronous changes within the terrestrial and marine realm at the triassic/jurassic boundary (csővár section, hungary). Rev. Palaeobot. Palynol. 156, 401–409 (2009).Article 

Google Scholar 
Hochuli, P. A., Hermann, E., Vigran, J. O., Bucher, H. & Weissert, H. Rapid demise and recovery of plant ecosystems across the end-permian extinction event. Global Planet. Change 74, 144–155 (2010).Article 
ADS 

Google Scholar 
Bonis, N. et al.Palaeoenvironmental changes and vegetation history during the Triassic-Jurassic transition (LPP Contribution Series No. 29, 2010).Bonis, N. R. & Kürschner, W. M. Vegetation history, diversity patterns, and climate change across the triassic/jurassic boundary. Paleobiology 38, 240–264 (2012).Article 

Google Scholar 
Visscher, H. et al. Environmental mutagenesis during the end-permian ecological crisis. Proc. Natl. Acad. Sci. 101, 12952–12956 (2004).Article 
ADS 
CAS 

Google Scholar  More

Ouellet, V. et al. Sci. Total Environ. 736, 139679 (2020).Article 
CAS 
PubMed 

Google Scholar 
Sutadian, A. D., Muttil, N., Yilmaz, A. G. & Perera, B. J. C. Environ. Monit. Assess. 188, 58 (2016).Article 
PubMed 

Google Scholar 
Murdoch, P. S., Baron, J. S. & Miller, T. L. J. Am. Water Resour. Assoc. 36, 347–366 (2000).Article 
CAS 

Google Scholar 
Hannah, D. M. & Garner, G. Prog Phys Geogr. 39, 68–92 (2015).Article 

Google Scholar 
Abbott, B. W. et al. Nat. Geosci. 12, 533–540 (2019).Article 
CAS 

Google Scholar 
Grill, G. et al. Nature 569, 215–221 (2019).Article 
CAS 
PubMed 

Google Scholar 
Hermanson, L. et al. Bull. Am. Meteorol. Soc. 103, E1117–E1129 (2022).Article 

Google Scholar 
Webb, B. W., Hannah, D. M., Moore, R. D., Brown, L. E. & Nobilis, F. Hydrol. Process. 22, 902–918 (2008).Article 

Google Scholar 
Hester, E. T. & Doyle, M. W. J. Am. Water Resour. Assoc. 47, 571–587 (2011).Article 

Google Scholar 
Schliemann, S. A., Grevstad, N. & Brazeau, R. H. Hydrol. Process 35, e14001 (2021).Article 

Google Scholar 
Jackson, F. L., Fryer, R. J., Hannah, D. M., Millar, C. P. & Malcolm, I. A. Sci. Total Environ. 612, 1543–1558 (2018).Article 
CAS 
PubMed 

Google Scholar 
O’Sullivan, A. M., Devito, K. J. & Curry, R. A. Catena 177, 70–83 (2019).Article 

Google Scholar 
Chang, H. & Psaris, M. Sci. Total Environ. 461, 587–600 (2013).Article 
PubMed 

Google Scholar 
Hester, E. T. & Bauman, K. S. J. Am. Water Resour. Assoc. 49, 328–342 (2013).Article 

Google Scholar 
Croghan, D., Van Loon, A. F., Sadler, J. P., Bradley, C. & Hannah, D. M. Hydrol. Process. 33, 144–159 (2018).Article 

Google Scholar 
Levia, D. F. et al. Nat. Geosci. 13, 656–658 (2020).Article 
CAS 

Google Scholar 
Nelson, K. C. & Palmer, M. A. J. Am. Water Resour. Assoc 43, 440–452 (2007).Article 

Google Scholar 
Heggenes, J. et al. River Res. Appl. 37, 743–765 (2021).Article 

Google Scholar 
Menberg, K., Blum, P., Kurylyk, B. L. & Bayer, P. Hydrol. Earth Syst. Sci. 18, 4453–4466 (2014).Article 

Google Scholar 
Tissen, C., Benz, S. A., Menberg, K., Bayer, P. & Blum, P. Environ. Res. Lett. 14, 104012 (2019).Article 
CAS 

Google Scholar 
Hannah, D. M. et al. Hydrol. Process. 36, e14525 (2022).Article 

Google Scholar 
Carothers, C. et al. Ecol. Soc. https://doi.org/10.5751/ES-11972-260116 (2021).Dugdale, S. J., Hannah, D. M. & Malcolm, I. A. Earth Sci. Rev. 175, 97–113 (2017).Article 

Google Scholar 
Wanders, N., van Vliet, M. T. H., Wada, Y., Bierkens, M. F. P. & van Beek, L. P. H. Water Resour. Res. 55, 2760–2778 (2019).Article 

Google Scholar 
Tavares, M. H. et al. Remote Sens. Environ. 241, 11172 (2020).Article 

Google Scholar 
Dugdale, S. J., Klaus, J. & Hannah, D. M. Water Resour. Res. 58, e2021WR031168 (2022).Article 

Google Scholar 
Mao, F. et al. Environ. Sci. Technol. 54, 9145–9158 (2020).Article 
CAS 
PubMed 

Google Scholar 
Hannah, D. M. et al. Hydrol. Process. 25, 1191–1200 (2011).Article 

Google Scholar 
Do, H. X., Gudmundsson, L., Leonard, M. & Westra, S. Earth Syst. Sci. Data 10, 765–785 (2018).Article 

Google Scholar  More

Mooney, H. et al. Biodiversity, climate change, and ecosystem services. Curr. Opin. Environ. Sustain. 1, 46–54 (2009).Article 

Google Scholar 
Perrings, C., Duraiappah, A., Larigauderie, A. & Mooney, H. The biodiversity and ecosystem services science-policy interface. Science 331, 1139–1140 (2011).Article 
ADS 
CAS 

Google Scholar 
Yachi, S. & Loreau, M. Biodiversity and ecosystem productivity in a fluctuating environment: The insurance hypothesis. Proc. Natl. Acad. Sci. USA 96, 1463–1468 (1999).Article 
ADS 
CAS 

Google Scholar 
Elmqvist, T. et al. Response diversity, ecosystem change, and resilience. Front. Ecol. Environ. 1, 488–494 (2003).Article 

Google Scholar 
Gonzalez, A. & Loreau, M. The causes and consequences of compensatory dynamics in ecological communities. Annu. Rev. Ecol. Evol. Syst. 40, 393–414 (2009).Article 

Google Scholar 
Blüthgen, N. & Klein, A.-M. Functional complementarity and specialisation: The role of biodiversity in plant–pollinator interactions. Basic Appl. Ecol. 12, 282–291 (2011).Article 

Google Scholar 
Brittain, C., Kremen, C. & Klein, A. M. Biodiversity buffers pollination from changes in environmental conditions. Glob. Change Biol. 19, 540–547 (2013).Article 
ADS 

Google Scholar 
Rader, R., Reilly, J., Bartomeus, I. & Winfree, R. Native bees buffer the negative impact of climate warming on honey bee pollination of watermelon crops. Glob. Chang. Biol. 19, 3103–3110 (2013).Article 
ADS 

Google Scholar 
Rogers, S. R., Tarpy, D. R. & Burrack, H. J. Bee species diversity enhances productivity and stability in a perennial crop. PLoS ONE 9, e97307 (2014).Article 
ADS 

Google Scholar 
Kühsel, S. & Blüthgen, N. High diversity stabilizes the thermal resilience of pollinator communities in intensively managed grasslands. Nat. Commun. 6, 1–10 (2015).Article 

Google Scholar 
Knop, E. et al. Rush hours in flower visitors over a day-night cycle. Insect Conserv. Divers. 11, 267–275 (2018).Article 

Google Scholar 
Goodwin, E. K., Rader, R., Encinas-Viso, F. & Saunders, M. E. Weather conditions affect the visitation frequency, richness and detectability of insect flower visitors in the Australian Alpine zone. Environ. Entomol. 50, 348–358 (2021).Article 

Google Scholar 
Feit, B. et al. Landscape complexity promotes resilience of biological pest control to climate change. Proc. Biol. Sci. 288, 20210547 (2021).
Google Scholar 
Tomas, F., Martínez-Crego, B., Hernán, G. & Santos, R. Responses of seagrass to anthropogenic and natural disturbances do not equally translate to its consumers. Glob. Chang. Biol. 21, 4021–4030 (2015).Article 
ADS 

Google Scholar 
Mori, A. S., Furukawa, T. & Sasaki, T. Response diversity determines the resilience of ecosystems to environmental change. Biol. Rev. 88, 349–364 (2013).Article 

Google Scholar 
Cariveau, D. P., Williams, N. M., Benjamin, F. E. & Winfree, R. Response diversity to land use occurs but does not consistently stabilise ecosystem services provided by native pollinators. Ecol. Lett. 16, 903–911 (2013).Article 

Google Scholar 
Garibaldi, L. A. et al. Wild pollinators enhance fruit set of crops regardless of honey bee abundance. Science 339, 1608. https://doi.org/10.1126/science.1230200 (2013).Article 
ADS 
CAS 

Google Scholar 
Kennedy, C. M. et al. A global quantitative synthesis of local and landscape effects on wild bee pollinators in agroecosystems. Ecol. Lett. 16, 584–599 (2013).Article 

Google Scholar 
Rader, R. et al. Non-bee insects are important contributors to global crop pollination. Proc. Natl. Acad. Sci. 113, 146–151 (2016).Article 
ADS 
CAS 

Google Scholar 
Klein, A.-M. et al. Importance of pollinators in changing landscapes for world crops. Proc. R. Soc. B Biol. Sci. 274, 303–313 (2007).Article 

Google Scholar 
Smith, M. R., Singh, G. M., Mozaffarian, D. & Myers, S. S. Effects of decreases of animal pollinators on human nutrition and global health: A modelling analysis. Lancet 386, 1964–1972 (2015).Article 

Google Scholar 
González-Varo, J. P. et al. Combined effects of global change pressures on animal-mediated pollination. Trends Ecol. Evol. 28, 524–530 (2013).Article 

Google Scholar 
Marshall, L. et al. The interplay of climate and land use change affects the distribution of EU bumblebees. Glob. Change Biol. 24, 101–116 (2018).Article 
ADS 

Google Scholar 
Millard, J. et al. Global effects of land-use intensity on local pollinator biodiversity. Nat. Commun. 12, 1–11 (2021).Article 
ADS 

Google Scholar 
Vasiliev, D. & Greenwood, S. The role of climate change in pollinator decline across the Northern Hemisphere is underestimated. Sci. Total Environ. 775, 145788 (2021).Article 
ADS 
CAS 

Google Scholar 
Steffan-Dewenter, I., Münzenberg, U., Bürger, C., Thies, C. & Tscharntke, T. Scale-dependent effects of landscape context on three pollinator guilds. Ecology 83, 1421–1432 (2002).Article 

Google Scholar 
Hass, A. L. et al. Landscape configurational heterogeneity by small-scale agriculture, not crop diversity, maintains pollinators and plant reproduction in western Europe. Proc. R. Soc. B Biol. Sci. https://doi.org/10.1098/rspb.2017.2242 (2018).Article 

Google Scholar 
Winfree, R. & Kremen, C. Are ecosystem services stabilized by differences among species? A test using crop pollination. Proc. R. Soc. B Biol. Sci. 276, 229–237 (2009).Article 

Google Scholar 
Jauker, F., Diekoetter, T., Schwarzbach, F. & Wolters, V. Pollinator dispersal in an agricultural matrix: Opposing responses of wild bees and hoverflies to landscape structure and distance from main habitat. Landsc. Ecol. 24, 547–555 (2009).Article 

Google Scholar 
Weiner, C. N., Werner, M., Linsenmair, K. E. & Blüthgen, N. Land-use impacts on plant–pollinator networks: Interaction strength and specialization predict pollinator declines. Ecology 95, 466–474 (2014).Article 

Google Scholar 
Chain-Guadarrama, A., Martínez-Salinas, A., Aristizábal, N. & Ricketts, T. H. Ecosystem services by birds and bees to coffee in a changing climate: A review of coffee berry borer control and pollination. Agric. Ecosyst. Environ. 280, 53–67 (2019).Article 

Google Scholar 
Hegland, S. J., Nielsen, A., Lázaro, A., Bjerknes, A. L. & Totland, Ø. How does climate warming affect plant–pollinator interactions?. Ecol. Lett. 12, 184–195 (2009).Article 

Google Scholar 
Bartomeus, I. et al. Contribution of insect pollinators to crop yield and quality varies with agricultural intensification. PeerJ 2, e328 (2014).Article 

Google Scholar 
Albrecht, M., Schmid, B., Hautier, Y. & Müller, C. B. Diverse pollinator communities enhance plant reproductive success. Proc. R. Soc. B Biol. Sci. 279, 4845–4852 (2012).Article 

Google Scholar 
Ellis, C. R., Feltham, H., Park, K., Hanley, N. & Goulson, D. Seasonal complementary in pollinators of soft-fruit crops. Basic Appl. Ecol. 19, 45–55 (2017).Article 

Google Scholar 
Brittain, C., Williams, N., Kremen, C. & Klein, A.-M. Synergistic effects of non-Apis bees and honey bees for pollination services. Proc. R. Soc. B Biol. Sci. 280, 20122767 (2013).Article 

Google Scholar 
Miñarro, M. & Twizell, K. W. Pollination services provided by wild insects to kiwifruit (Actinidia deliciosa). Apidologie 46, 276–285 (2015).Article 

Google Scholar 
Senapathi, D., Goddard, M. A., Kunin, W. E. & Baldock, K. C. Landscape impacts on pollinator communities in temperate systems: Evidence and knowledge gaps. Funct. Ecol. 31, 26–37 (2017).Article 

Google Scholar 
Papanikolaou, A. D., Kuehn, I., Frenzel, M. & Schweiger, O. Landscape heterogeneity enhances stability of wild bee abundance under highly varying temperature, but not under highly varying precipitation. Landsc. Ecol. 32, 581–593 (2017).Article 

Google Scholar 
Papanikolaou, A. D., Kühn, I., Frenzel, M. & Schweiger, O. Semi-natural habitats mitigate the effects of temperature rise on wild bees. J. Appl. Ecol. 54, 527–536 (2017).Article 

Google Scholar 
Orford, K. A., Vaughan, I. P. & Memmott, J. The forgotten flies: The importance of non-syrphid Diptera as pollinators. Proc. R. Soc. B Biol. Sci. 282, 20142934 (2015).Article 

Google Scholar 
Settele, J., Bishop, J. & Potts, S. G. Climate change impacts on pollination. Nat. Plants 2, 1–3 (2016).Article 

Google Scholar 
Taki, H., Okabe, K., Makino, S. I., Yamaura, Y. & Sueyoshi, M. Contribution of small insects to pollination of common buckwheat, a distylous crop. Ann. Appl. Biol. 155, 121–129 (2009).Article 

Google Scholar 
Krkošková, B. & Mrazova, Z. Prophylactic components of buckwheat. Food Res. Int. 38, 561–568 (2005).Article 

Google Scholar 
Campbell, J. W., Irvin, A., Irvin, H., Stanley-Stahr, C. & Ellis, J. D. Insect visitors to flowering buckwheat, Fagopyrum esculentum (Polygonales: Polygonaceae), in north-central Florida. Fla. Entomol. 99, 264–268 (2016).Article 

Google Scholar 
Hadley, N. F. Water Relations of Terrestrial Arthropods (CUP Archive, 1994).
Google Scholar 
Sgolastra, F. et al. Temporal activity patterns in a flower visitor community of Dictamnus albus in relation to some biotic and abiotic factors. Bull. Insectol. 69, 291–300 (2016).
Google Scholar 
Vicens, N. & Bosch, J. Weather-dependent pollinator activity in an apple orchard, with special reference to Osmia cornuta and Apis mellifera (Hymenoptera: Megachilidae and Apidae). Environ. Entomol. 29, 413–420 (2000).Article 

Google Scholar 
Carlucci, M. B., Brancalion, P. H., Rodrigues, R. R., Loyola, R. & Cianciaruso, M. V. Functional traits and ecosystem services in ecological restoration. Restor. Ecol. 28, 1372–1383 (2020).Article 

Google Scholar 
Lavorel, S. Plant functional effects on ecosystem services. (2013).Defra. (ed Food and Rural Affairs Department for Environment) (2019).Agency, J. M. Amedas, https://tenki.jp/past/2019/09/amedas/ (2019).Jacquemart, A.-L., Gillet, C. & Cawoy, V. Floral visitors and the importance of honey bee on buckwheat (Fagopyrum esculentum Moench) in central Belgium. J. Hortic. Sci. Biotechnol. 82, 104–108 (2007).Article 

Google Scholar 
Taki, H. et al. Effects of landscape metrics on Apis and non-Apis pollinators and seed set in common buckwheat. Basic Appl. Ecol. 11, 594–602 (2010).Article 

Google Scholar 
Dray, S., Legendre, P. & Peres-Neto, P. R. Spatial modelling: A comprehensive framework for principal coordinate analysis of neighbour matrices (PCNM). Ecol. Model. 196, 483–493 (2006).Article 

Google Scholar 
Legendre, P. & Legendre, L. Numerical Ecology (Elsevier, 2012).MATH 

Google Scholar 
Dray S, et al. adespatial: Multivariate Multiscale Spatial Analysis. R package version 0.3-20, https://CRAN.R-project.org/package=adespatial. (2022).Benjamin, F. E., Reilly, J. R. & Winfree, R. Pollinator body size mediates the scale at which land use drives crop pollination services. J. Appl. Ecol. 51, 440–449 (2014).Article 

Google Scholar 
Földesi, R. et al. Relationships between wild bees, hoverflies and pollination success in apple orchards with different landscape contexts. Agric. For. Entomol. 18, 68–75 (2016).Article 

Google Scholar 
Oksanen J, et al. vegan: Community Ecology Package. R package version 2.6-4. https://CRAN.R-project.org/package=vegan. (2022)Bürkner, P.-C. brms: An R package for Bayesian multilevel models using Stan. J. Stat. Softw. 80, 1–28 (2017).Article 

Google Scholar 
Gelman, A., Carlin, J. B., Stern, H. S. & Rubin, D. B. Bayesian Data Analysis (Chapman and Hall/CRC, 1995).Book 
MATH 

Google Scholar 
Team, R. C. R: A Language and Environment for Statistical Computing (2019).Sasaki, H. & Wagatsuma, T. Bumblebees (Apidae: Hymenoptera) are the main pollinators of common buckwheat, Fogopyrum esculentum, in Hokkaido, Japan. Appl. Entomol. Zool. 42, 659–661 (2007).Article 

Google Scholar 
Nagano, Y., Miyashita, T., Taki, H. & Yokoi, T. Diversity of co-flowering plants at field margins potentially sustains an abundance of insects visiting buckwheat, Fagopyrum esculentum, in an agricultural landscape. Ecol. Res. 36, 882–891 (2021).Article 

Google Scholar 
Samra, S., Samocha, Y., Eisikowitch, D. & Vaknin, Y. Can ants equal honeybees as effective pollinators of the energy crop Jatropha curcas L. under Mediterranean conditions?. Gcb Bioenergy 6, 756–767 (2014).Article 

Google Scholar 
Sugiura, N., Miyazaki, S. & Nagaishi, S. A supplementary contribution of ants in the pollination of an orchid, Epipactis thunbergii, usually pollinated by hover flies. Plant Syst. Evol. 258, 17–26 (2006).Article 

Google Scholar 
Natsume, K., Hayashi, S. & Miyashita, T. Ants are effective pollinators of common buckwheat Fagopyrum esculentum. Agric. For. Entomol. 24, 446–452 (2022).Article 

Google Scholar 
Carvalheiro, L. G., Seymour, C. L., Nicolson, S. W. & Veldtman, R. Creating patches of native flowers facilitates crop pollination in large agricultural fields: Mango as a case study. J. Appl. Ecol. 49, 1373–1383 (2012).Article 

Google Scholar 
Michiyama, H., Arikuni, M. & Hirano, T. Effect of air temperature on the growth, flowering and ripening in common buckwheat. In The Procceeding of the 8th ISB (2001)Isbell, F. et al. High plant diversity is needed to maintain ecosystem services. Nature 477, 199-U196. https://doi.org/10.1038/nature10282 (2011).Article 
ADS 
CAS 

Google Scholar 
McCain, C. M. & Colwell, R. K. Assessing the threat to montane biodiversity from discordant shifts in temperature and precipitation in a changing climate. Ecol. Lett. 14, 1236–1245 (2011).Article 

Google Scholar 
Choi, S.-W. Effects of weather factors on the abundance and diversity of moths in a temperate deciduous mixed forest of Korea. Zool. Sci. 25, 53–58 (2008).Article 

Google Scholar 
Feldmeier, S. et al. Climate versus weather extremes: Temporal predictor resolution matters for future rather than current regional species distribution models. Divers. Distrib. 24, 1047–1060 (2018).Article 

Google Scholar  More

Strains belonging to the same species display distinct growth dynamics on the marine polysaccharide alginateWe first quantified the growth dynamics of the 12 Vibrionaceae strains (Supplementary Table 1) on alginate in well-mixed batch cultures. Growth of populations was initiated at approximately the same inoculum density (105 colony forming units (c.f.u.) ml−1). We tracked the growth dynamics by measuring the optical density at 600 nm and compared the maximum population size reached over the course of 36 h (Fig. 1 and S1). We found significant differences in the maximal optical density achieved by different strains within each species (Fig. 1 and S1). In V. splendidus, strains 12B01 and FF6 reached a lower maximum population size compared to strains 1S124 and 13B01 (Fig. 1 and S1A). In V. cyclitrophicus, strain ZF270 reached a lower maximum population size compared to strains 1F175, 1F111, and ZF28 (Fig. 1 and S1A). Similarly, in V. sp. F13, strain 9ZC77 reached a lower maximum population size than strains 9CS106, 9ZC13, and ZF57 (Fig. 1 and S1A). These findings suggest that some strains are limited in their growth abilities in well-mixed environments, perhaps as a consequence of differences in the amount and activity of enzymes they release (Supplementary Table 1).Fig. 1: Vibrionaceae strains differ in their growth dynamics on the marine polysaccharide alginate under well-mixed conditions.Maximum optical density (measured at 600 nm) achieved by populations of strains belonging to Vibrio splendidus, Vibrio cyclitrophicus, and Vibrio sp. F13 during the course of a 36 h growth cycle on the same concentration (0.1% weight/volume) of the polysaccharide alginate. Points and error bars indicate the mean of measurements across populations within each ecotype (npopulations = 3) and the 95% confidence interval (CI), respectively. Different letters indicate statistically significant differences between strains within one species (One-way ANOVA and Dunnett’s post-hoc test; V. splendidus: p  More

Pawlowski, J. et al. CBOL Protist working group: barcoding eukaryotic richness beyond the animal, plant, and fungal kingdoms. PLOS Biol. 10, e1001419 (2012).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
del Campo, J. et al. The others: our biased perspective of eukaryotic genomes. Trends Ecol. Evol. 29, 252–259 (2014).Article 
PubMed 
PubMed Central 

Google Scholar 
Handbook of the Protists (Springer, 2017). https://doi.org/10.1007/978-3-319-28149-0.Lang, B. F., O’Kelly, C., Nerad, T., Gray, M. W. & Burger, G. The closest unicellular relatives of animals. Curr. Biol. 12, 1773–1778 (2002).Article 
CAS 
PubMed 

Google Scholar 
del Campo, J. et al. Ecological and evolutionary significance of novel protist lineages. Eur. J. Protistol. 55, 4–11 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Grau-Bové, X. et al. Dynamics of genomic innovation in the unicellular ancestry of animals. Life 6, e26036 (2017).
Google Scholar 
Gawryluk, R. M. R. et al. Non-photosynthetic predators are sister to red algae. Nature 572, 240–243 (2019).Article 
CAS 
PubMed 

Google Scholar 
Gabr, A., Grossman, A. R. & Bhattacharya, D. Paulinella, a model for understanding plastid primary endosymbiosis. J. Phycol. 56, 837–843 (2020).Article 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Gao, Z., Karlsson, I., Geisen, S., Kowalchuk, G. & Jousset, A. Protists: Puppet masters of the rhizosphere microbiome. Trends Plant Sci. 24, 165–176 (2019).Article 
CAS 
PubMed 

Google Scholar 
Caron, D. A. New accomplishments and approaches for assessing protistan diversity and ecology in natural ecosystems. Bioscience 59, 287–299 (2009).Article 

Google Scholar 
Gooday, A. J., Schoenle, A., Dolan, J. R. & Arndt, H. Protist diversity and function in the dark ocean: Challenging the paradigms of deep-sea ecology with special emphasis on foraminiferans and naked protists. Eur. J. Protistol. 75, 125721 (2020).Article 
PubMed 

Google Scholar 
Stoecker, D. K., Johnson, M. D., de Vargas, C. & Not, F. Acquired phototrophy in aquatic protists. Aquat. Microb. Ecol. 57, 279–310 (2009).Article 

Google Scholar 
Strom, S. L., Benner, R., Ziegler, S. & Dagg, M. J. Planktonic grazers are a potentially important source of marine dissolved organic carbon. Limnol. Oceanogr. 42, 1364–1374 (1997).Article 
ADS 
CAS 

Google Scholar 
Orsi, W. D. et al. Identifying protist consumers of photosynthetic picoeukaryotes in the surface ocean using stable isotope probing. Environ. Microbiol. 20, 815–827 (2018).Article 
CAS 
PubMed 

Google Scholar 
Corno, G. & Jürgens, K. Direct and indirect effects of protist predation on population size structure of a bacterial strain with high phenotypic plasticity. Appl. Environ. Microbiol. 72, 78–86 (2006).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Mahé, F. et al. Parasites dominate hyperdiverse soil protist communities in Neotropical rainforests. Nat. Ecol. Evol. 1, 91 (2017).Article 
PubMed 

Google Scholar 
Ruppert, K. M., Kline, R. J. & Rahman, M. S. Past, present, and future perspectives of environmental DNA (eDNA) metabarcoding: A systematic review in methods, monitoring, and applications of global eDNA. Glob. Ecol. Conserv. 17, e00547 (2019).Article 

Google Scholar 
Epstein, S. & López-García, P. “Missing” protists: a molecular prospective. Biodivers. Conserv. 17, 261–276 (2008).Article 

Google Scholar 
López-García, P., Rodríguez-Valera, F., Pedrós-Alió, C. & Moreira, D. Unexpected diversity of small eukaryotes in deep-sea Antarctic plankton. Nature 409, 603–607 (2001).Article 
ADS 
PubMed 

Google Scholar 
Lovejoy, C., Massana, R. & Pedrós-Alió, C. Diversity and distribution of marine microbial eukaryotes in the Arctic Ocean and adjacent seas. Appl. Environ. Microbiol. 72, 3085–3095 (2006).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Worden, A. Z., Cuvelier, M. L. & Bartlett, D. H. In-depth analyses of marine microbial community genomics. Trends Microbiol. 14, 331–336 (2006).Article 
CAS 
PubMed 

Google Scholar 
Countway, P. D. et al. Distinct protistan assemblages characterize the euphotic zone and deep sea (2500 m) of the western North Atlantic (Sargasso Sea and Gulf Stream). Environ. Microbiol. 9, 1219–1232 (2007).Article 
CAS 
PubMed 

Google Scholar 
Massana, R. & Pedrós-Alió, C. Unveiling new microbial eukaryotes in the surface ocean. Curr. Opin. Microbiol. 11, 213–218 (2008).Article 
PubMed 

Google Scholar 
Alexander, E. et al. Microbial eukaryotes in the hypersaline anoxic L’Atalante deep-sea basin. Environ. Microbiol. 11, 360–381 (2009).Article 
CAS 
PubMed 

Google Scholar 
Stoeck, T. et al. Multiple marker parallel tag environmental DNA sequencing reveals a highly complex eukaryotic community in marine anoxic water. Mol. Ecol. 19, 21–31 (2010).Article 
CAS 
PubMed 

Google Scholar 
Logares, R. et al. Patterns of rare and abundant marine microbial eukaryotes. Curr. Biol. 24, 813–821 (2014).Article 
CAS 
PubMed 

Google Scholar 
de Vargas, C. et al. Eukaryotic plankton diversity in the sunlit ocean. Science 348, 150 (2015).Article 

Google Scholar 
Fell, J. W., Scorzetti, G., Connell, L. & Craig, S. Biodiversity of micro-eukaryotes in Antarctic Dry Valley soils with More

Darwin, C. The Descent of Man and Selection in Relation to Sex. (John Murray, 1871).Andersson, M. Sexual Selection. (Princeton University Press, 1994).Shuster, S. & Wade, M. J. Mating Systems and Strategies. (Princeton University Press, 2003).Gosden, T. P. & Svensson, E. I. Spatial and temporal dynamics in a sexual selection mosaic. Evolution 62, 845–856 (2008).Article 
PubMed 

Google Scholar 
Kasumovic, M. M., Bruce, M. J., Andrade, M. C. B. & Herberstein, M. E. Spatial and temporal demographic variation drives within-season fluctuations in sexual selection. Evolution 62, 2316–2325 (2008).Article 
PubMed 

Google Scholar 
Mobley, K. B. & Jones, A. G. Environmental, demographic, and genetic mating system variation among five geographically distinct dusky pipefish (Syngnathus floridae) populations. Mol. Ecol. 18, 1476–1490 (2009).Article 
PubMed 

Google Scholar 
Hoffer, J. N., Mariën, J., Ellers, J. & Koene, J. M. Sexual selection gradients change over time in a simultaneous hermaphrodite. eLife 6, e25139 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
Sih, A., Montiglio, P.-O., Wey, T. W. & Fogarty, S. Altered physical and social conditions produce rapidly reversible mating systems in water striders. Behav. Ecol. 28, 632–639 (2017).Article 

Google Scholar 
Preston, B. T., Stevenson, I. R., Pemberton, J. M. & Wilson, K. Dominant rams lose out by sperm depletion. Nature 409, 681–682 (2001).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Cornwallis, C. K. & Uller, T. Towards an evolutionary ecology of sexual traits. Trends Ecol. Evol. 25, 145–152 (2010).Article 
PubMed 

Google Scholar 
Forsgren, E., Amundsen, T., Borg, A. A. & Bjelvenmark, J. Unusually dynamic sex roles in a fish. Nature 429, 551–554 (2004).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Hare, R. M. & Simmons, L. W. Sexual selection maintains a female-specific character in a species with dynamic sex roles. Behav. Ecol. 32, 609–616 (2021).Article 

Google Scholar 
Fox, R. J., Donelson, J. M., Schunter, C., Ravasi, T. & Gaitán-Espitia, J. D. Beyond buying time: the role of plasticity in phenotypic adaptation to rapid environmental change. Philos. Trans. R. Soc. B 374, 20180174 (2019).Article 

Google Scholar 
Ingleby, F. C., Hunt, J. & Hosken, D. J. The role of genotype-by-environment interactions in sexual selection. J. Evol. Biol. 23, 2031–2045 (2010).Article 
CAS 
PubMed 

Google Scholar 
Lindström, J., Pike, T. W., Blount, J. D. & Metcalfe, N. B. Optimization of resource allocation can explain the temporal dynamics and honesty of sexual signals. Am. Nat. 174, 515–525 (2009).Article 
PubMed 

Google Scholar 
Janicke, T., David, P. & Chapuis, E. Environment-dependent sexual selection: Bateman’s parameters under varying levels of food availability. Am. Nat. 185, 756–768 (2015).Article 
PubMed 

Google Scholar 
Morimoto, J., Pizzari, T. & Wigby, S. Developmental environment effects on sexual selection in male and female Drosophila melanogaster. PLoS ONE 11, e0154468 (2016).Article 
PubMed 
PubMed Central 

Google Scholar 
Cattelan, S., Evans, J. P., Garcia-Gonzalez, F., Morbiato, E. & Pilastro, A. Dietary stress increases the total opportunity for sexual selection and modifies selection on condition-dependent traits. Ecol. Lett. 23, 447–456 (2020).Article 
PubMed 

Google Scholar 
Glavaschi, A., Cattelan, S., Grapputo, A. & Pilastro, A. Imminent risk of predation reduces the relative strength of postcopulatory sexual selection in the guppy. Philos. Trans. R. Soc. B 375, 20200076 (2020).Article 

Google Scholar 
Clark, D. C., DeBano, S. J. & Moore, A. J. The influence of environmental quality on sexual selection in Nauphoeta cinerea (Dictyoptera: Blaberidae). Behav. Ecol. 8, 46–53 (1997).Article 

Google Scholar 
Emlen, S. & Oring, L. Ecology, sexual selection and the evolution of mating systems. Science 197, 215–223 (1977).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Liker, A., Freckleton, R. P. & Székely, T. The evolution of sex roles in birds is related to adult sex ratio. Nat. Commun. 4, 1–6 (2013).Article 

Google Scholar 
Wacker, S. et al. Operational sex ratio but not density affects sexual selection in a fish. Evolution 67, 1937–1949 (2013).Article 
PubMed 

Google Scholar 
Wacker, S., Ness, M. H., Östlund-Nilsson, S. & Amundsen, T. Social structure affects mating competition in a damselfish. Coral Reefs 36, 1279–1289 (2017).Article 
ADS 

Google Scholar 
Janicke, T. & Morrow, E. H. Operational sex ratio predicts the opportunity and direction of sexual selection across animals. Ecol. Lett. 21, 384–391 (2018).Article 
PubMed 

Google Scholar 
Procter, D. S., Moore, A. J. & Miller, C. W. The form of sexual selection arising from male-male competition depends on the presence of females in the social environment. J. Evol. Biol. 25, 803–812 (2012).Article 
CAS 
PubMed 

Google Scholar 
Eldakar, O. T., Dlugos, M. J., Pepper, J. W. & Wilson, D. S. Population structure mediates sexual conflict in Water striders. Science 326, 816–816 (2009).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Martin, A. M., Festa-Bianchet, M., Coltman, D. W. & Pelletier, F. Demographic drivers of age-dependent sexual selection. J. Evol. Biol. 29, 1437–1446 (2016).Article 
CAS 
PubMed 

Google Scholar 
Pilakouta, N. & Ålund, M. Sexual selection and environmental change: what do we know and what comes next? Curr. Zool. 67, 293–298 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Kahn, A. T., Dolstra, T., Jennions, M. D. & Backwell, P. R. Y. Strategic male courtship effort varies in concert with adaptive shifts in female mating preferences. Behav. Ecol. 24, 906–913 (2013).Article 

Google Scholar 
Jordan, L. A. & Brooks, R. C. Recent social history alters male courtship preferences. Evolution 66, 280–287 (2012).Article 
PubMed 

Google Scholar 
Wilson, D. R., Nelson, X. J. & Evans, C. S. Seizing the opportunity: Subordinate male fowl respond rapidly to variation in social context. Ethology 115, 996–1004 (2009).Article 

Google Scholar 
Gwynne, D. T., Bailey, W. J. & Annells, A. The sex in short supply for matings varies over small Spatial scales in a Katydid (Kawanaphila nartee, Orthoptera: Tettigoniidae). Behav. Ecol. Sociobiol. 42, 157–162 (1998).Article 

Google Scholar 
Fedina, T. Y. & Lewis, S. M. Female mate choice across mating stages and between sequential mates in flour beetles. J. Evol. Biol. 20, 2138–2143 (2007).Article 
CAS 
PubMed 

Google Scholar 
Clark, H. L. & Backwell, P. R. Y. Temporal and spatial variation in female mating preferences in a fiddler crab. Behav. Ecol. Sociobiol. 69, 1779–1784 (2015).Article 

Google Scholar 
Serbezov, D., Bernatchez, L., Olsen, E. M. & Vøllestad, L. A. Mating patterns and determinants of individual reproductive success in brown trout (Salmo trutta) revealed by parentage analysis of an entire stream living population. Mol. Ecol. 19, 3193–3205 (2010).Article 
PubMed 

Google Scholar 
Gerlach, N. M., McGlothlin, J. W., Parker, P. G. & Ketterson, E. D. Reinterpreting Bateman gradients: multiple mating and selection in both sexes of a songbird species. Behav. Ecol. 23, 1078–1088 (2012).Article 

Google Scholar 
Dubuc, C., Ruiz-Lambides, A. & Widdig, A. Variance in male lifetime reproductive success and estimation of the degree of polygyny in a primate. Behav. Ecol. 25, 878–889 (2014).Article 
PubMed 
PubMed Central 

Google Scholar 
Breuer, T. et al. Variance in the male reproductive success of western gorillas: acquiring females is just the beginning. Behav. Ecol. Sociobiol. 64, 515–528 (2010).Article 

Google Scholar 
Germain, R. R., Hallworth, M. T., Kaiser, S. A., Sillett, T. S. & Webster, M. S. Variance in within-pair reproductive success influences the opportunity for selection annually and over the lifetimes of males in a multi-brooded songbird. Evolution 75, 915–930 (2021).Article 
PubMed 

Google Scholar 
Lande, R. & Arnold, S. J. The measurement of selection on correlated characters. Evolution 37, 1210–1226 (1983).Article 
PubMed 

Google Scholar 
Klug, H., Heuschele, J., Jennions, M. D. & Kokko, H. The mismeasurement of sexual selection. J. Evol. Biol. 23, 447–462 (2010).Article 
CAS 
PubMed 

Google Scholar 
Jennions, M. D., Kokko, H. & Klug, H. The opportunity to be misled in studies of sexual selection. J. Evol. Biol. 25, 591–598 (2012).Article 
CAS 
PubMed 

Google Scholar 
Krakauer, A. H., Webster, M. S., Duval, E. H., Jones, A. G. & Shuster, S. M. The opportunity for sexual selection: not mismeasured, just misunderstood. J. Evol. Biol. 24, 2064–2071 (2011).Article 
CAS 
PubMed 

Google Scholar 
Hebets, E. A., Stafstrom, J. A., Rodriguez, R. L. & Wilgers, D. J. Enigmatic ornamentation eases male reliance on courtship performance for mating success. Anim. Behav. 81, 963–972 (2011).Article 

Google Scholar 
Fitzpatrick, J. L. & Lüpold, S. Sexual selection and the evolution of sperm quality. Mol. Hum. Reprod. 20, 1180–1189 (2014).Article 
PubMed 

Google Scholar 
Jones, A. G. On the opportunity for sexual selection, the Bateman gradient and the maximum intensity of sexual selection. Evolution 63, 1673–1684 (2009).Article 
PubMed 

Google Scholar 
Henshaw, J. M., Kahn, A. T. & Fritzsche, K. A rigorous comparison of sexual selection indexes via simulations of diverse mating systems. Proc. Natl Acad. Sci. USA 113, E300–E308 (2016).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
Evans, J. P. & Garcia-Gonzalez, F. The total opportunity for sexual selection and the integration of pre- and post-mating episodes of sexual selection in a complex world. J. Evol. Biol. 29, 2338–2361 (2016).Article 
CAS 
PubMed 

Google Scholar 
Downhower, J. F., Blumer, L. S. & Brown, L. Opportunity for selection: an appropriate measure for evaluating variation in the potential for selection? Evolution 41, 1395–1400 (1987).Article 
PubMed 

Google Scholar 
Klug, H. & Stone, L. More than just noise: Chance, mating success, and sexual selection. Ecol. Evol. 11, 6326–6340 (2021).Article 
PubMed 
PubMed Central 

Google Scholar 
Anthes, N., Häderer, I. K., Michiels, N. K. & Janicke, T. Measuring and interpreting sexual selection metrics: evaluation and guidelines. Methods Ecol. Evol. 8, 918–931 (2016).Article 

Google Scholar 
Klug, H., Lindström, K. & Kokko, H. Who to include in measures of sexual selection is no trivial matter. Ecol. Lett. 13, 1094–1102 (2010).Article 
PubMed 

Google Scholar 
Collet, J. M., Dean, R. F., Worley, K., Richardson, D. S. & Pizzari, T. The measure and significance of Bateman’s principles. Proc. R. Soc. B 281, 20132973 (2014).Article 
PubMed 
PubMed Central 

Google Scholar 
Collet, J., Richardson, D. S., Worley, K. & Pizzari, T. Sexual selection and the differential effect of polyandry. Proc. Natl Acad. Sci. USA 109, 8641–8645 (2012).Article 
ADS 
CAS 
PubMed 
PubMed Central 

Google Scholar 
McDonald, G. C., Spurgin, L. G., Fairfield, E. A., Richardson, D. S. & Pizzari, T. Pre- and postcopulatory sexual selection favor aggressive, young males in polyandrous groups of red junglefowl. Evolution 71, 1653–1669 (2017).Article 
PubMed 

Google Scholar 
Morimoto, J. et al. Sex peptide receptor-regulated polyandry modulates the balance of pre- and post-copulatory sexual selection in Drosophila. Nat. Commun. 10, 283 (2019).Article 
ADS 
PubMed 
PubMed Central 

Google Scholar 
Shuster, S. M., Willen, R. M., Keane, B. & Solomon, N. G. Alternative mating tactics in socially monogamous prairie voles, Microtus ochrogaster. Front. Ecol. Evol. 7, 7 (2019).Article 

Google Scholar 
Dowling, J. & Webster, M. S. Working with what you’ve got: unattractive males show greater mate-guarding effort in a duetting songbird. Biol. Lett. 13, 20160682 (2017).Article 
PubMed 
PubMed Central 

Google Scholar 
Pizzari, T. & McDonald, G. C. Sexual selection in socially structured, polyandrous populations: Some insights from the fowl. Adv. Study Behav. 51, 77–141 (2019).Article 

Google Scholar 
Archer, M. S. & Elgar, M. A. Female preference for multiple partners: sperm competition in the hide beetle, Dermestes maculatus (DeGeer). Anim. Behav. 58, 669–675 (1999).Article 
CAS 
PubMed 

Google Scholar 
Qvarnström, A. & Forsgren, E. Should females prefer dominant males? Trends Ecol. Evol. 13, 498–501 (1998).Article 
PubMed 

Google Scholar 
Webster, M. S., Tarvin, K. A., Tuttle, E. M. & Pruett-Jones, S. Promiscuity drives sexual selection in a socially monogamous bird. Evolution 61, 2205–2211 (2007).Article 
PubMed 

Google Scholar 
Brunton, D. H. Energy expenditure in reproductive effort of male and female Killdeer (Charadrius vociferus). Auk 105, 553–564 (1988).Article 

Google Scholar 
Johnson, L. S., Hicks, B. G. & Masters, B. S. Increased cuckoldry as a cost of breeding late for male house wrens (Troglodytes aedon). Behav. Ecol. 13, 670–675 (2002).Article 

Google Scholar 
Boinski, S. Mating patterns in squirrel monkeys (Saimiri oerstedi): implications for seasonal sexual dimorphism. Behav. Ecol. Sociobiol. 21, 13–21 (1987).Article 

Google Scholar 
McDonald, G. C., Spurgin, L. G., Fairfield, E. A., Richardson, D. S. & Pizzari, T. Differential female sociality is linked with the fine-scale structure of sexual interactions in replicate groups of red junglefowl, Gallus gallus. Proc. R. Soc. B 286, 20191734 (2019).Article 
PubMed 
PubMed Central 

Google Scholar 
Carleial, R. et al. Temporal dynamics of competitive fertilization in social groups of red junglefowl (Gallus gallus) shed new light on avian sperm competition. Philos. Trans. R. Soc. B 375, 20200081 (2020).Article 

Google Scholar 
Lessells, C. M. & Birkhead, T. R. Mechanisms of sperm competition in birds: mathematical models. Behav. Ecol. Sociobiol. 27, 325–337 (1990).Article 

Google Scholar 
Taborsky, T., Oliveira, R. F. & Brockmann, H. J. The Evolution of Alternative Reproductive Tactics: Concepts and Questions. in Alternative Reproductive Tactics: An Integrative Approach (Cambridge University Press, 2008).Ghislandi, P. G. et al. Resource availability, mating opportunity and sexual selection intensity influence the expression of male alternative reproductive tactics. J. Evol. Biol. 31, 1035–1046 (2018).Article 
PubMed 

Google Scholar 
Lehtonen, T. K., Wong, B. B. M. & Lindström, K. Fluctuating mate preferences in a marine fish. Biol. Lett. 6, 21–23 (2010).Article 
PubMed 

Google Scholar 
Chaine, A. S. & Lyon, B. E. Adaptive plasticity in female mate choice dampens sexual selection on male ornaments in the lark bunting. Science 319, 459–462 (2008).Article 
ADS 
CAS 
PubMed 

Google Scholar 
R Core Team. R: A Language and Environment for Statistical Computing. (R Foundation for Statistical Computing, 2019).Bates, D., Mächler, M., Bolker, B. & Walker, S. Fitting Linear Mixed-Effects Models Using lme4. J. Stat. Softw. 67, 1–48 (2015).Article 

Google Scholar 
Kuznetsova, A., Brockhoff, P. B. & Christensen, R. H. lmerTest package: tests in linear mixed effects models. J. Stat. Softw. 82, 1–26 (2017).Article 

Google Scholar 
Oklander, L. I., Kowalewski, M. & Corach, D. Male reproductive strategies in black and gold howler monkeys (Alouatta caraya). Am. J. Primatol. 76, 43–55 (2014).Article 
PubMed 

Google Scholar 
Pröhl, H. & Hödl, W. Parental investment, potential reproductive rates, and mating system in the strawberry dart-poison frog, Dendrobates pumilio. Behav. Ecol. Sociobiol. 46, 215–220 (1999).Article 

Google Scholar 
Turnell, B. R. & Shaw, K. L. High opportunity for postcopulatory sexual selection under field conditions. Evolution 69, 2094–2104 (2015).Article 
PubMed 

Google Scholar 
Gill, L. F., van Schaik, J., von Bayern, A. M. P. & Gahr, M. L. Genetic monogamy despite frequent extrapair copulations in “strictly monogamous” wild jackdaws. Behav. Ecol. 31, 247–260 (2020).Article 
PubMed 

Google Scholar 
Carleial, R., McDonald, G. C. & Pizzari, T. Dynamic phenotypic correlates of social status and mating effort in male and female red junglefowl, Gallus gallus. J. Evol. Biol. 33, 22–40 (2020).Article 
PubMed 

Google Scholar 
McDonald, G. C. & Pizzari, T. Structure of sexual networks determines the operation of sexual selection. Proc. Natl Acad. Sci. USA 115, E53–E61 (2018).Article 
ADS 
CAS 
PubMed 

Google Scholar 
Janicke, T., Häderer, I. K., Lajeunesse, M. J. & Anthes, N. Darwinian sex roles confirmed across the animal kingdom. Sci. Adv. 2, e1500983 (2016).Article 
ADS 
PubMed 
PubMed Central 

Google Scholar 
Webster, M. S., Pruett-Jones, S., Westneat, D. F. & Arnold, S. J. Measuring the effects of pairing success, extra-pair copulations and mate quality on the opportunity for sexual selection. Evolution 49, 1147–1157 (1995).PubMed 

Google Scholar 
Etches, R. J. Reproduction in Poultry. (CABI, 1996).Schielzeth, H. Simple means to improve the interpretability of regression coefficients: Interpretation of regression coefficients. Methods Ecol. Evol. 1, 103–113 (2010).Article 

Google Scholar 
Løvlie, H., Cornwallis, C. K. & Pizzari, T. Male mounting alone reduces female promiscuity in the fowl. Curr. Biol. 15, 1222–1227 (2005).Article 
PubMed 

Google Scholar 
Berglund, A. Many mates make male pipefish choosy. Behaviour 132, 213–218 (1995).Article 

Google Scholar 
Carleial, R., Pizzari, T., Richardson, D. S. & McDonald, G. C. Data for: Disentangling the causes of temporal variation in the opportunity for sexual selection. figshare Dataset (2023) https://doi.org/10.6084/m9.figshare.21902133.v1.McLain, D. K., Burnette, L. B. & Deeds, D. A. Within season variation in the intensity of sexual selection on body size in the bug Margus obscurator (Hemiptera Coreidae). Ethol. Ecol. Evol. 5, 75–86 (1993).Article 

Google Scholar 
Schlicht, E. & Kempenaers, B. Effects of social and extra-pair mating on sexual selection in Blue tits (Cyanistes caeruleus). Evolution 67, 1420–1434 (2013).PubMed 

Google Scholar  More