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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2025.1514216</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Brief Research Report</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>First spawning record of the widespread Indo-Pacific <italic>Pavona maldivensis</italic> observed in the Red Sea</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Garcias-Bonet</surname>
<given-names>Neus</given-names>
</name>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Casartelli</surname>
<given-names>Marco</given-names>
</name>
<uri xlink:href="https://loop.frontiersin.org/people/2649530"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Vimercati</surname>
<given-names>Silvia</given-names>
</name>
<uri xlink:href="https://loop.frontiersin.org/people/2314317"/>
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<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Benzoni</surname>
<given-names>Francesca</given-names>
</name>
<uri xlink:href="https://loop.frontiersin.org/people/1363019"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Peixoto</surname>
<given-names>Raquel S.</given-names>
</name>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/122390"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
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</contrib-group>
<aff id="aff1">
<institution>Biological and Environmental Sciences and Engineering Division (BESE), King Abdullah University of Science and Technology (KAUST)</institution>, <addr-line>Thuwal</addr-line>, <country>Saudi Arabia</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Masaya Morita, University of the Ryukyus, Japan</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Saki Harii, University of the Ryukyus, Japan</p>
<p>Go Suzuki, Seikai National Fisheries Research Institute, Japan</p>
<p>Emmeline Jamodiong, University of the Ryukyus, Japan</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Neus Garcias-Bonet, <email xlink:href="mailto:neus.garciasbonet@kaust.edu.sa">neus.garciasbonet@kaust.edu.sa</email>; Raquel S. Peixoto, <email xlink:href="mailto:raquel.peixoto@kaust.edu.sa">raquel.peixoto@kaust.edu.sa</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>25</day>
<month>02</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>12</volume>
<elocation-id>1514216</elocation-id>
<history>
<date date-type="received">
<day>20</day>
<month>10</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>06</day>
<month>02</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Garcias-Bonet, Casartelli, Vimercati, Benzoni and Peixoto</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Garcias-Bonet, Casartelli, Vimercati, Benzoni and Peixoto</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Information on coral reproductive biology and coral spawning times is crucial to advance basic and applied research and inform strategies for coral reef conservation and restoration. Important efforts have been made to collate coral spawning times and reproductive patterns in global and regional datasets. Here, we report and document the first <italic>in situ</italic> spawning of <italic>Pavona maldivensis</italic> Gardiner, 1905, observed in the Red Sea. A medium size colony was observed releasing sperm on the full moon night on 23 May 2024, at sunset time. Our observations suggest that the widespread Indo-Pacific <italic>P. maldivensis</italic> is likely gonochoric. This first report on the <italic>in situ</italic> spawning timing for <italic>P. maldivensis</italic> contributes to expanding coral spawning databases and provides valuable data on its reproductive biology, which is relevant for coral restoration and conservation efforts.</p>
</abstract>
<kwd-group>
<kwd>coral reproductive biology</kwd>
<kwd>coral gametes</kwd>
<kwd>broadcast spawners</kwd>
<kwd>coral sperm release</kwd>
<kwd>Red Sea</kwd>
</kwd-group>    <contract-num rid="cn001">FCC/1/1973-51- 01, REI/1/4984-01-01, BAS/1/1095-01-01</contract-num>    <contract-sponsor id="cn001">King Abdullah University of Science and Technology<named-content content-type="fundref-id">10.13039/501100004052</named-content>
</contract-sponsor>
<counts>
<fig-count count="1"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="25"/>
<page-count count="4"/>
<word-count count="1150"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Discoveries</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<title>Introduction</title>
<p>The current global coral crisis has pushed forward the development of a wide range of strategies to improve coral resilience to withstand the pressing anthropogenic impacts (<xref ref-type="bibr" rid="B25">Voolstra et&#xa0;al., 2021</xref>). Some of these interventions rely on successfully collecting gametes to sexually propagate corals. Therefore, detailed information on coral spawning is crucial to advance basic and applied research and inform strategies for coral reef conservation and restoration. Currently, reproduction data is available for over 300 scleractinian species in 61 genera in the Indo-Pacific (<xref ref-type="bibr" rid="B9">Harrison, 2011</xref>; <xref ref-type="bibr" rid="B2">Baird et&#xa0;al., 2021</xref>), where over 600 hard coral species are reported (<xref ref-type="bibr" rid="B24">Veron et&#xa0;al., 2015</xref>). In the Red Sea, where research on coral reproduction historically started in the Gulf of Aqaba (<xref ref-type="bibr" rid="B17">Rinkevich and Loya, 1979</xref>; <xref ref-type="bibr" rid="B21">Shlesinger and Loya, 1985</xref>; <xref ref-type="bibr" rid="B20">Shlesinger et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B16">Rapuano et&#xa0;al., 2017</xref>), synchronous spawning of scleractinian corals at central latitudes has been observed around full moon nights in spring (<xref ref-type="bibr" rid="B4">Bouwmeester et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B14">Osman et&#xa0;al., 2024</xref>). In the Indo-Pacific region, most spawning observations are on species of the genus <italic>Acropora</italic>, with 38% of the total spawning records, while information on other coral genera is relatively scarce (<xref ref-type="bibr" rid="B2">Baird et&#xa0;al., 2021</xref>).</p>
<p>
<italic>Pavona</italic> Lamarck, 1801, is commonly observed in shallow Indo-Pacific coral reefs, including the Red Sea (<xref ref-type="bibr" rid="B22">Terraneo et&#xa0;al., 2017</xref>), but few studies have described its reproductive biology. Species of <italic>Pavona</italic> are almost exclusively gonochoric and broadcast spawners (<xref ref-type="bibr" rid="B20">Shlesinger et&#xa0;al., 1998</xref>; <xref ref-type="bibr" rid="B9">Harrison, 2011</xref>). <italic>In situ</italic> spawning has been reported two and three days after the full moon of November in <italic>Pavona cactus</italic> Forsk&#xe5;l, 1775, in the Great Barrier Reef (<xref ref-type="bibr" rid="B11">Marshall and Stephenson, 1933</xref>), of March in <italic>Pavona gigantea</italic> Verrill, 1869, in the Galapagos Islands, Ecuador (<xref ref-type="bibr" rid="B6">Glynn et&#xa0;al., 1996</xref>), from January to April for <italic>Pavona varians</italic> Verrill, 1864, and <italic>Pavona</italic> sp. (later formally described as <italic>Pavona chiriquiensis</italic> Glynn, Mat&#xe9; &amp; Stemann, 2001) in the Gulf of Chiriqu&#xed;, Panama (<xref ref-type="bibr" rid="B8">Glynn et&#xa0;al., 2000</xref>), and in October for <italic>Pavona clavus</italic> Dana, 1846, in Contadora Island, Panama (<xref ref-type="bibr" rid="B7">Glynn et&#xa0;al., 2011</xref>). <italic>In situ</italic> spawning was observed six days after full moon of August in <italic>Pavona decussata</italic> Dana, 1846, in Kochi Prefecture, Japan (<xref ref-type="bibr" rid="B12">Mezaki et&#xa0;al., 2014</xref>). In the Red Sea, no <italic>in situ</italic> spawning observations have been reported so far for <italic>Pavona</italic>; however, April has been inferred as the spawning month in <italic>P. varians</italic>, based on the oocyte maturity (<xref ref-type="bibr" rid="B4">Bouwmeester et&#xa0;al., 2015</xref>). Here, we report the first <italic>in situ</italic> spawning observation of <italic>Pavona maldivensis</italic> Gardiner, 1905.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<title>Materials and methods</title>
<p>In April and May 2024, <italic>in situ</italic> surveys to document coral spawning were conducted in Al Fahal Reef (22&#xb0;18&#x2019;21&#x201d; N; 38&#xb0;57&#x2019;44&#x201d; E), a midshore reef located in the central Red Sea (<xref ref-type="bibr" rid="B5">Garcias-Bonet et&#xa0;al., 2024</xref>). Underwater surveys took place by SCUBA divers during two consecutive months on 21&#x2013;25 April 2024 and 20&#x2013;24 May 2024, covering the full moon nights on 24 April and 23 May 2024. Surveys were conducted from 17:30 to 19:00 and from 21:30 to 23:00 at depths from 8&#x2013;10 meters using red light torches to minimize the negative impact of light on coral spawning. Spawning observations and timing were noted and pictures and videos were taken with an Olympus TG-6 camera equipped with an Olympus PT-059 underwater housing (OM System). <italic>In situ</italic> seawater temperature was monitored using a Multiparameter CTD probe (Ocean Seven 310, Idronaut) as part of a sustained environmental data observation platform (<xref ref-type="bibr" rid="B5">Garcias-Bonet et&#xa0;al., 2024</xref>).</p>
</sec>
<sec id="s3" sec-type="results">
<title>Results and discussion</title>
<p>We report and document the first <italic>in situ</italic> spawning observation of <italic>Pavona maldivensis</italic> Gardiner, 1905 (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>) in the central Red Sea (Al Fahal Reef) on the full moon night, on 23 May 2024 at 18:49h, ten minutes before sunset time (18:59h), when <italic>in situ</italic> daily mean seawater temperature was 29.68&#xb0;C. A medium size <italic>P. maldivensis</italic> colony (80 cm in diameter) at a depth of 9 m was observed releasing sperm in a dense cloud surrounding the entire colony for seven minutes (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>, <xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Movie 1</bold>
</xref>). No eggs seemed to be released from the colony, suggesting that <italic>P. maldivensis</italic> is likely gonochoric, in agreement with reproductive data available for other <italic>Pavona</italic> species. For instance, <italic>Pavona varians</italic> has been reported as gonochoric in the Red Sea (<xref ref-type="bibr" rid="B4">Bouwmeester et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B20">Shlesinger et&#xa0;al., 1998</xref>) and mostly gonochoric with some hermaphrodite colonies in Eastern Pacific (<xref ref-type="bibr" rid="B8">Glynn et&#xa0;al., 2000</xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>
<bold>(A)</bold> Picture of a <italic>Pavona maldivensis</italic> colony in the Red Sea. Image by Francesca Benzoni. <bold>(B)</bold> Detail of a <italic>P. maldivensis</italic> colony <italic>in vivo</italic>. The colony exhibits the characteristic knob-shaped structure, with corallites defined by the species&#x2019; distinct raised, rounded, and well-defined wall. The corallite septa are alternating in height and thickness. Image by Marco Casartelli. <bold>(C)</bold> <italic>In situ</italic> spawning observation, with a dense cloud of sperm being released (arrows), from <italic>P. maldivensis</italic> in Al Fahal Reef in central Red Sea. Image by Neus Garcias-Bonet.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1514216-g001.tif"/>
</fig>
<p>Contrary to the spawning synchrony observed in many scleractinian corals (<xref ref-type="bibr" rid="B3">Baird et&#xa0;al., 2009</xref>, <xref ref-type="bibr" rid="B2">2021</xref>), spawning times in <italic>Pavona</italic> genus doesn&#x2019;t seem to be synchronized, with <italic>in situ</italic> observations ranging from midday to few hours before sunrise around full moon nights across different months. Similarly to our spawning observation at sunset time in <italic>P. maldivensis, Pavona</italic> sp. spawning was reported shortly after sunset in Panam&#xe1; (<xref ref-type="bibr" rid="B8">Glynn et&#xa0;al., 2000</xref>), <italic>P. gigantea</italic> spawning was reported in the late afternoon (two hours before sunset) in Galapagos Islands, Ecuador (<xref ref-type="bibr" rid="B6">Glynn et&#xa0;al., 1996</xref>), and the release of eggs in <italic>Pavona explanulata</italic> was observed three hours after sunset in Taiwan (<xref ref-type="bibr" rid="B10">Lin and Nozawa, 2017</xref>). Contrary, <italic>Pavona</italic> sp. spawning was reported at midday in Thailand (<xref ref-type="bibr" rid="B15">Plathong et&#xa0;al., 2006</xref>), male and female colonies of <italic>P. varians</italic> were observed spawning two hours before sunrise in Panam&#xe1; (<xref ref-type="bibr" rid="B8">Glynn et&#xa0;al., 2000</xref>) and <italic>P. decussata</italic> was observed spawning about one hour before sunrise in Japan (<xref ref-type="bibr" rid="B12">Mezaki et&#xa0;al., 2014</xref>).</p>
<p>The spawning observation of <italic>P. maldivensis</italic> reported here is based on a single male colony; therefore, further research, increasing the observation time window and number of monitored colonies, is needed to fully describe the spawning patterns, characterize the reproductive biology and confirm the gonochorism of <italic>P. maldivensis</italic> in the Red Sea. <italic>P. maldivensis</italic> is known to be widespread from the Red Sea to the East Pacific Region (<xref ref-type="bibr" rid="B19">Sheppard, 1991</xref>; <xref ref-type="bibr" rid="B23">Veron, 2000</xref>; <xref ref-type="bibr" rid="B18">Salvat et&#xa0;al., 2016</xref>). In the Red Sea, it can be commonly observed from very shallow to upper mesophotic low-light environments such as under reef crest ledges and underhangs (<xref ref-type="bibr" rid="B1">Al Tawaha et&#xa0;al., 2019</xref>), with <italic>Pavona</italic> genus accounting on average for less than 1% of the benthic cover in the central Red Sea (<xref ref-type="bibr" rid="B13">Monroe et&#xa0;al., 2018</xref>). This first report on the <italic>in situ</italic> spawning timing for <italic>P. maldivensis</italic> in the Red Sea contributes to expanding global coral spawning databases and provides valuable data on its reproductive biology, which is relevant for coral restoration and conservation efforts.</p>
</sec>
</body>
<back>
<sec id="s4" sec-type="data-availability">
<title>Data availability statement</title>
<p>The original contributions presented in the study are included in the article/<xref ref-type="supplementary-material" rid="SF1">
<bold>Supplementary Material</bold>
</xref>. Further inquiries can be directed to the corresponding author/s.</p>
</sec>
<sec id="s5" sec-type="ethics-statement">
<title>Ethics statement</title>
<p>The manuscript presents research on animals that do not require ethical approval for their study.</p>
</sec>
<sec id="s6" sec-type="author-contributions">
<title>Author contributions</title>
<p>NG-B: Conceptualization, Methodology, Writing &#x2013; original draft. MC: Conceptualization, Methodology, Writing &#x2013; review &amp; editing. SV: Methodology, Writing &#x2013; review &amp; editing. FB: Conceptualization, Writing &#x2013; review &amp; editing. RP: Conceptualization, Funding acquisition, Writing &#x2013; review &amp; editing.</p>
</sec>
<sec id="s7" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research, authorship, and/or publication of this article. NG-B and RP acknowledge funding from KAUST (grants FCC/1/1973-51- 01, REI/1/4984-01-01 and BAS/1/1095-01-01).</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>We would like to thank the Coastal and Marine Resources Core Lab team at KAUST.</p>
</ack>
<sec id="s8" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec id="s9" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
</sec>
<sec id="s10" sec-type="disclaimer">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec id="s11" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2025.1514216/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2025.1514216/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Video1.mov" id="SF1" mimetype="video/quicktime">
<label>Supplementary Movie 1</label>
<caption>
<p>
<italic>In situ</italic> spawning observation from a <italic>Pavona maldivensis</italic> colony in Al Fahal Reef in central Red Sea on the full moon night in May, on the 23rd of May 2024, after sunset time (18:50h). File format: Apple QuickTime Movie (MOV).</p>
</caption>
</supplementary-material>
</sec>
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