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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.2024.1402044</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Marine Science</subject>
<subj-group>
<subject>Review</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Updating and validating seagrass ecosystem knowledge in the gulf of California: a comprehensive review</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Ram&#xed;rez-Z&#xfa;&#xf1;iga</surname>
<given-names>Magali Alejandra</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
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<xref ref-type="aff" rid="aff2">
<sup>2</sup>
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<contrib contrib-type="author">
<name>
<surname>P&#xe9;rez-Estrada</surname>
<given-names>Claudia Jeannette</given-names>
</name>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
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<contrib contrib-type="author">
<name>
<surname>L&#xf3;pez-Calder&#xf3;n</surname>
<given-names>Jorge Manuel</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Cannon</surname>
<given-names>Abigail Libbin</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
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<contrib contrib-type="author">
<name>
<surname>Vanderplank</surname>
<given-names>Sula</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<xref ref-type="aff" rid="aff7">
<sup>7</sup>
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<contrib contrib-type="author" corresp="yes">
<name>
<surname>Favoretto</surname>
<given-names>Fabio</given-names>
</name>
<xref ref-type="aff" rid="aff8">
<sup>8</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
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<aff id="aff1">
<sup>1</sup>
<institution>Universidad Aut&#xf3;noma de Baja California Sur, Departamento Acad&#xe9;mico de Ciencias Marinas y Costeras</institution>, <addr-line>La Paz, BCS</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Centro para la Biodiversidad Marina y la Conservaci&#xf3;n, A.C.</institution>, <addr-line>La Paz, BCS</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>Centro de Investigaciones Biol&#xf3;gicas del Noroeste S.C. Departamento de Ecolog&#xed;a Pesquera</institution>, <addr-line>La Paz, BCS</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Universidad Aut&#xf3;noma de Baja California, Facultad de Ciencias Marinas</institution>, <addr-line>Ensenada, BC</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Department of Life Sciences, Texas A&amp;M University</institution>, <addr-line>Corpus Christi, TX</addr-line>, <country>United States</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Centro de Estudios del Medio Ambiente, Eco-Alianza de Loreto</institution>, <addr-line>Loreto, BCS</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff7">
<sup>7</sup>
<institution>Conserva Loreto, The Ocean Foundation</institution>, <addr-line>Loreto, BCS</addr-line>, <country>Mexico</country>
</aff>
<aff id="aff8">
<sup>8</sup>
<institution>Scripps Institution of Oceanography, University of California San Diego</institution>, <addr-line>La Jolla, CA</addr-line>, <country>United States</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Lorenzo Mari, Polytechnic University of Milan, Italy</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Tania Cota, Center for Research and Advanced Studies - M&#xe9;rida Unit, Mexico</p>
<p>Susan Bell, University of South Florida, United States</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Fabio Favoretto, <email xlink:href="mailto:ffavoretto@ucsd.edu">ffavoretto@ucsd.edu</email>
</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>17</day>
<month>05</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1402044</elocation-id>
<history>
<date date-type="received">
<day>16</day>
<month>03</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>24</day>
<month>04</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Ram&#xed;rez-Z&#xfa;&#xf1;iga, P&#xe9;rez-Estrada, L&#xf3;pez-Calder&#xf3;n, Cannon, Vanderplank and Favoretto</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Ram&#xed;rez-Z&#xfa;&#xf1;iga, P&#xe9;rez-Estrada, L&#xf3;pez-Calder&#xf3;n, Cannon, Vanderplank and Favoretto</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>Seagrasses are globally acknowledged as crucial habitats as they provide a variety of ecosystem services. Mexico&#x2019;s legislation protects most of these marine plants; however, the protection often fails in application. The Gulf of California, despite being a biodiversity hotspot, has scant data on seagrasses. Here, human activity and climate change increasingly threaten these coastal ecosystems, with conservation and research efforts lacking coordination at a regional level. Our manuscript aimed to review and standardize existing data on Gulf of California seagrass species, ensuring open access for data updates; pinpointing conservation deficiencies; and guiding future research. We have added new records to the official public data, but we were able to recapture only 25% of the seagrass locations meaning a potential reduction in their historical distribution of 45.8%. Even though Mexico&#x2019;s legislation protects some species of seagrasses, it protection often fails in the application. We identified that only 6.1% of the seagrass locations are within protected areas that recognize their presence in their management plans (e.g., the Balandra Flora and Fauna Protected Area and the Upper Gulf of California and Colorado River Delta Biosphere Reserve). At least 55.9% of seagrass records are associated with potentially damaging activities like pollution, coastal modification or biological resources use, while 23% are exposed to higher frequency of marine heatwaves. Given the importance of seagrass meadows under Mexican law and their internationally recognized ecological value, sharing current information and guiding research is essential. Our study seeks to galvanize renewed research initiatives and raise more awareness on the conservation of the Gulf of California&#x2019;s seagrasses.</p>
</abstract>
<kwd-group>
<kwd>critical habitat</kwd>
<kwd>blue carbon</kwd>
<kwd>spatial monitoring</kwd>
<kwd>habitat conservation</kwd>
<kwd>Mexico</kwd>
<kwd>Eastern Pacific</kwd>
</kwd-group>
<counts>
<fig-count count="4"/>
<table-count count="0"/>
<equation-count count="0"/>
<ref-count count="119"/>
<page-count count="12"/>
<word-count count="5694"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Marine Conservation and Sustainability</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Seagrasses provide essential ecosystem services including primary productivity, nutrient cycling, habitat provision, coastal protection, and carbon sequestration (<xref ref-type="bibr" rid="B17">Costanza et&#xa0;al., 1997</xref>; <xref ref-type="bibr" rid="B75">Ondiviela et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B110">Unsworth et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B109">United Nations Environment Programme, 2020</xref>). However, these vital meadows are in global decline due to climate change, overfishing, coastal development, and pollution (<xref ref-type="bibr" rid="B78">Orth et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B115">Waycott et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B103">Short et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B10">Brodie and de Ramon N&#x2019;yeurt, 2018</xref>). The extent of seagrass loss is not fully known, largely because of insufficient historical data on their distribution (<xref ref-type="bibr" rid="B10">Brodie and de Ramon N&#x2019;yeurt, 2018</xref>; <xref ref-type="bibr" rid="B64">McKenzie et&#xa0;al., 2020</xref>). This lack of data is pronounced in the Gulf of California, where there has been a historical deficit in mapping and research of seagrass species (<xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B64">McKenzie et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B29">Dunic et&#xa0;al., 2021</xref>).</p>
<p>The Gulf of California, along with the Pacific coast of the Baja California Peninsula, contains significant stretches of seagrasses within the Mexican Pacific (<xref ref-type="bibr" rid="B108">UNEP-WCMC and Short, 2021</xref>). Four seagrass species have been documented in the Gulf of California; three are protected under Mexican law (NOM-059-SEMARNAT-2010, <xref ref-type="bibr" rid="B27">DOF, 2019</xref>). &#x2014;<italic>Zostera marina</italic> L. 1753, <italic>Halophila decipiens</italic> Ostenfeld 1902 (both under Special Protection), and <italic>Halodule wrightii</italic> Ascherson 1868 (considered Threatened). The fourth species, <italic>Ruppia maritima</italic> L. 1753, is not included in this legislation. Although widely distributed, the most prominent seagrass meadows are found in lagoons and bays, which makes them vulnerable to anthropogenic impacts (<xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>, <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>). Given the importance of seagrasses for their benefits to human well-being, as blue carbon sinks and indirect contributors to climate change mitigation (<xref ref-type="bibr" rid="B42">Friess, 2023</xref>), it is crucial to gain a thorough understanding of their conditions and establish a research baseline. To date, the absence of coordinated regional research has resulted in a fragmented comprehension of seagrass ecosystems, and there is a significant lack of large-scale mapping, monitoring, or analysis of spatial trends. Consequently, the available official data on seagrass distribution in the Gulf of California is incomplete, inconsistent, or not current (e.g., UNEP-WCMC). Since these data are utilized by stakeholders and policymakers both locally and internationally, the need for accuracy is critical.</p>
<p>This review aims to 1) provide a comprehensive, validated, open-source dataset of seagrass locations in the Gulf of California by compiling an extensive collection of published studies, gray literature, herbarium records, and input from experts; 2) identify current research and conservation shortcomings; and 3) establish a baseline to direct future scientific endeavors. This effort is expected to act as a catalyst for new studies and support informed conservation tactics for seagrasses in the Gulf of California.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study area</title>
<p>The Gulf of California extends over 1,200 km, encompassing 40 coastal lagoons and 922 islands, and is part of the unique Cortezian marine ecoregion (<xref ref-type="bibr" rid="B104">Spalding et&#xa0;al., 2007</xref>). The regional climate is influenced by the El Ni&#xf1;o Southern Oscillation (ENSO) and the Pacific Decadal Oscillation (PDO), factors contributing to substantial oceanic changes (<xref ref-type="bibr" rid="B62">Luch-Cota et&#xa0;al., 2013</xref>). In recent decades, the Gulf of California has experienced pronounced warming, affecting a multitude of marine species and habitats (<xref ref-type="bibr" rid="B72">Nev&#xe1;rez-Mart&#xed;nez et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B114">Velarde et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B32">Favoretto et&#xa0;al., 2022</xref>; <xref ref-type="bibr" rid="B94">S&#xe1;nchez-Cabeza et&#xa0;al., 2022</xref>). It features distinct latitudinal gradients in sea surface temperature and chlorophyll-<italic>a</italic> concentrations, which delimit three primary oceanographic regions (<xref ref-type="bibr" rid="B11">Brusca, 2010</xref>; <xref ref-type="bibr" rid="B107">Ulate et&#xa0;al., 2016</xref>). These regions all experience similar maximum summer sea surface temperatures around 30&#xb0;C. However, during winter, the northern region records cooler temperatures (16&#xb0;C), the central region exhibits an intermediate average temperature of around 20&#xb0;C while the southern region remains warmer (22&#xb0;C). Also, the northern region has the highest annual average chlorophyll-<italic>a</italic> concentration at 1.6 mg m<sup>&#x2013;3</sup>, compared to the southern region, which has an average of 0.5 mg m<sup>&#x2013;3</sup> (<xref ref-type="bibr" rid="B107">Ulate et&#xa0;al., 2016</xref>).</p>
<p>Seagrass meadows, found in the Gulf of California&#x2019;s bays, lagoons, and inlets, are increasingly vulnerable to human activities. The eastern lagoons are heavily exploited for agriculture, aquaculture, settlement, and fishing. In contrast, the western bays and inlets face pressures from coastal development and mining activities (<xref ref-type="bibr" rid="B3">&#xc1;lvarez-Borrego, 1983</xref>; <xref ref-type="bibr" rid="B72">Nev&#xe1;rez-Mart&#xed;nez et&#xa0;al., 2001</xref>; <xref ref-type="bibr" rid="B114">Velarde et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B46">Gonz&#xe1;lez-Abraham et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B80">P&#xe1;ez-Osuna et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B94">S&#xe1;nchez-Cabeza et&#xa0;al., 2022</xref>). These cumulative pressures threaten the most favorable habitats for seagrass meadows, underscoring the urgency to address current and future anthropogenic threats in the Gulf of California.</p>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Systematic review protocol</title>
<p>In conducting this review, we undertook a thorough examination of academic and gray literature, national and international herbarium records, and floristic surveys. Our research employed &#x2018;Google Scholar&#x2019; for the English search terms &#x2018;Seagrasses&#x2019;, &#x2018;Gulf of California&#x2019; and the name of the species recorded in the region &#x2018;<italic>Zostera marina</italic>,&#x2019; <italic>&#x2018;Ruppia maritima</italic>,&#x2019; <italic>&#x2018;Halodule wrightii</italic>,&#x2019; <italic>&#x2018;Halophila decipiens&#x2019;</italic>. Equivalent Spanish terms were used, replacing &#x2018;Seagrasses&#x2019; with &#x2018;Pastos marinos&#x2019; and &#x2018;Gulf of California&#x2019; with &#x2018;Golfo de California.&#x2019; The search was conducted from July 2019 to March 2021. Additionally, we classified the studies according to their primary focus and depicted their temporal progression graphically. While our primary emphasis was on peer-reviewed scientific, we included other sources of information such as technical and governmental reports, theses at the undergraduate, master&#x2019;s, and doctoral levels due to their significant insights into the distribution and condition of the seagrass meadows.</p>
<p>Subsequently, we evaluated seven online herbarium collections from the following institutions: the University of Arizona (ARIZ), Arizona State University (ASU), Autonomous University of Baja California (CMMEX), National Herbarium of Mexico (MEXU) of the National Autonomous University of Mexico, Smithsonian National Museum of Natural History (SI NMNH), Jes&#xfa;s Gonz&#xe1;lez Ortega Herbarium at the Autonomous University of Sinaloa (UAS), and the University of Sonora (UNISON-USON). From these archives, we extracted details such as species identification, collection dates, locality, and geographical coordinates. Each entry was then organized by location, facilitating the creation of maps that illustrate the historical distribution of each seagrass species throughout the Gulf of California.</p>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Assessing seagrass distribution and potential stressors</title>
<sec id="s2_3_1">
<label>2.3.1</label>
<title>Validation and assessment of distribution decline</title>
<p>Historical distribution records were obtained through the systematic review. On the field, we executed aerial reconnaissance to authenticate the presence or absence of seagrasses at coordinates reported in historical records. Drone flights, with altitudes oscillating between 10m and 60m, were primarily employed for intertidal seagrass detection. In addition, snorkeling expeditions complemented these aerial inspections. We classified records as &#x201c;present&#x201d;, when the record was validated at historically reported coordinates, when seagrasses were not found, the record was classified as &#x201c;absent&#x201d;. Finally, where the historical record was conspicuously imprecise (e.g., coordinates fell inland) or when pixel classification and drone imagery were unattainable, records were designated as &#x201c;data deficient&#x201d;. The <xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 1</bold>
</xref> includes information about the tools used in each location, and specific data about them.</p>
</sec>
<sec id="s2_3_2">
<label>2.3.2</label>
<title>Assessment of protection level</title>
<p>We define the protection level as the existence of conservation measures for each record such as the presence of a protected area. The visualization of this data is available at <xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 2</bold>
</xref>. As the presence of a protected area is not indication of active protection on the seagrass ecosystem, we also reviewed management plans to discriminate the ones mentioning seagrasses explicitly or not. We then classified areas not protected as the ones outside any management polygon and separated protected areas according to their category established by the National Commission of Protected Natural Areas (CONANP) (e.g., National Park, Flora and Fauna protection area, Biosphere reserve, or Ramsar site). We used spatial data from government digital repositories (<xref ref-type="bibr" rid="B16">Comisi&#xf3;n Nacional de &#xc1;reas Naturales Protegidas, 2021</xref>) Natural Protected Area locations, utilizing Geographical Information Systems (QGIS, v. 3.18).</p>
</sec>
<sec id="s2_3_3">
<label>2.3.3</label>
<title>Assessment of potential human and climate stressors</title>
<p>We identified potential human stressors based on the presence of activities known to harm seagrass health and gauged climate stressors using marine heatwave frequency as a proxy. Due to the varied thermal tolerances of species and a lack of specific quantitative data, we propose that increased marine heatwave frequency, indicative of extreme climate events, suggests elevated ecosystem stress.</p>
<p>For human stressors, we consulted scientific literature, local reports, and protected area management plans from governmental archives. We noted the presence of activities such as agricultural runoff leading to eutrophication, coastal development activities like construction and dredging that destroy habitats, and harmful fishing practices. These stressors often co-occur, amplifying their impact. Spatial correlation to localities was conducted using QGIS.</p>
<p>Regarding climate stressors, we analyzed sea surface temperature (SST) data from the Reynolds&#x2019; OISST dataset, spanning 1982-2020, with a resolution of 0.25 x 0.25 degrees. Marine heatwaves, defined as periods where daily SSTs exceeded the 90th percentile threshold for at least five consecutive days, were identified and their frequency was modeled using a GLM with a Poisson distribution in R, utilizing the &#x201c;heatwaveR&#x201d; package (<xref ref-type="bibr" rid="B49">Hobday et al., 2016</xref>; <xref ref-type="bibr" rid="B100">Schlegel and Smit, 2018</xref>). Rates of change in heatwave frequency were calculated for each grid point and used as a proxy for potential future impacts, normalized on a 0-1 scale for compatibility with other criteria. Detailed methodologies and additional statistical treatments are provided in the <xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 2</bold>
</xref>.</p>
</sec>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Seagrass distribution</title>
<p>Our research compiled records of seagrass species from 28 locations in the Gulf of California, with 20 on the eastern coast and six on the Baja California Peninsula (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). We compared our results with the official data from Ocean Data Viewer where only nine seagrass locations have been reported (<xref ref-type="bibr" rid="B108">UNEP-WCMC &amp; Short, 2021</xref>). This data does not provide information about the extension or the name of the locality, only provide the coordinates. Additionally, they focus on only two of the four species of seagrass in the Gulf of California (<italic>Zostera marina</italic> and <italic>Halodule wrightii</italic>). Particularly in Los Cabos, this database includes a record of <italic>Phillospadix torreyi</italic> and <italic>Phillospadix scouleri</italic>, both species that are not recorded within the Gulf of California.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Dual maps compare seagrass distributions along the coast. <bold>(A)</bold> displays records obtained from UNEP-WCMC (2021), while <bold>(B)</bold> reflects this study&#x2019;s updated findings, showcasing expanded and refined seagrass locations. Data information with geographic coordinates is available in <xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 1</bold>
</xref>.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1402044-g001.tif"/>
</fig>
<p>We identified primary seagrass localities through record frequency, occurrence rates, and data on their distribution. First, The Canal del Infiernillo, between the Sonora coast and Tiburon Island, includes all bays and estuaries within, then Bah&#xed;a Concepci&#xf3;n, as a second critical seagrass habitat in the Gulf of California.</p>
<p>In this work, we include new records. Records prior to 2018 are observations by the authors that have not been previously published. While the records obtained during our field validation efforts were collected between 2018 and 2022 (<xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 1</bold>
</xref>). In 2020, we identified <italic>H. wrightii</italic> in Ensenada de la Paz channel. Additionally, <italic>H. decipiens</italic> was in San Gabriel Bay on Isla Espiritu Santo in 2018, and at the Pichilingue research unit of the Autonomous University of Baja California Sur in 2022.</p>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Human and climate stressors</title>
<p>The analysis of our results shows that several locations are likely facing multiple stress factors. 17 of 28 historical locations are subject to the three main types of human-induced stressors: pollution, coastal modification, and overuse of biological resources (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>). While climate stress occurs in 12 locations. However, this phenomenon is most pronounced at spots like the Colorado River delta, Bahia Concepci&#xf3;n, and El Verde coastal lagoon (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Spatial distribution of human stressors and marine heatwave frequencies over seagrass meadows. <bold>(A)</bold> illustrates a cumulative score for human-induced stressors impacting seagrass meadows, with values ranging from low (0.00) to high (1.00), depicted as a gradient of colors from green (low) to red (high). <bold>(B)</bold> represents the slope of marine heatwave (MHW) frequencies, calculated for each seagrass locality, with cooler colors indicating lower frequencies and warmer colors denoting higher frequencies. Both panels are geographically aligned to demonstrate the spatial correlation between human activities and thermal anomalies across the studied seagrass habitats.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1402044-g002.tif"/>
</fig>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Conservation and management</title>
<p>We observed that most of the seagrass records are located outside protected areas or in areas with low levels of protection that lack management strategies and plans, such as Ramsar sites. While 13.2% of the records are found within protected areas (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). However, its presence and relevance are only recognized in the management plan of the Balandra Flora and Fauna Protection Area (<xref ref-type="bibr" rid="B26">DOF, 2012</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Bar chart illustrating the distribution of seagrass records by level of protection.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1402044-g003.tif"/>
</fig>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Current information and knowledge gaps</title>
<p>Our comprehensive review yielded 77 documents related to seagrasses in the Gulf of California, classified into eight categories. 42 of them were &#x201c;Peer-reviewed&#x201d; academic articles. The categories of &#x201c;Books&#x201d; and &#x201c;Book Chapters&#x201d; followed, with 11 and 6 records respectively. Since 1996, ten theses have been registered spanning bachelor&#x2019;s, master&#x2019;s, and doctoral degrees, although only three have been published as peer-reviewed papers. Despite relatively limited, we included two outreach papers and five reports of projects of research in our analysis.</p>
<p>Although research into seagrasses began more than 100 years ago, the rise of peer-reviewed publications occurred at the end of the 20th century, in the 1980s, when the peak of research in a decade was reached, with 12 of them produced between 1980 and 1988. However, from 1990 to 2021, this trend reversed, adding a total of 25 peer-reviewed publications in 31 years, i.e., a rate of 0.80, less than one publication per year. Despite this slowdown, starting in 2010, an important moment stands out in the production of research on seagrasses, with the increase in theses and reports, indicating a resurgence of interest in research and the continued study of seagrasses in the region (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>).</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Seagrass research in the Gulf of California by <bold>(A)</bold> type of document and <bold>(B)</bold> research topics accumulated over the years.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1402044-g004.tif"/>
</fig>
<p>During the first years, there is a trend towards taxonomic and ethnobotanical studies. This period is characterized by no increase in the number of studies until the 1970s when studies focused on the distribution of seagrasses emerge, and the first research mentioning them indirectly (Indirect observations) appears.</p>
<p>Ecology and conservation are the most studied topics, increased almost constantly since the first publication in 1977 and 1992, respectively. Both topics reaching a maximum number of studies in 2021. The last decade of the 20th century was an important time for research, as new lines of research emerged, such as genetics, blue carbon, in which it is necessary to redouble efforts (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>).</p>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<sec id="s4_1">
<label>4.1</label>
<title>Seagrass distribution</title>
<p>The mapping of seagrass habitats remains a significant challenge, despite recent advances in technology that promise to narrow the information gap. Even the wide extent of seagrasses in the Gulf of California, our validation data results only confirmed the presence of seagrasses at 13 out of the 28 historically noted localities (<xref ref-type="supplementary-material" rid="SM1">
<bold>Datasheet 1</bold>
</xref>). This indicates a potential decline of 46.4% from their known historical distribution, a concerning trend also reported by <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al. (2016)</xref>.</p>
<p>The seagrass meadows of the Canal del Infiernillo are the most important within the Gulf of California, due they cover the largest spatial area and appear to be the most consistent in their distribution over time. It extends between the Sonora coast and Tiburon Island, includes all bays and estuaries therein. The Canal del Infiernillo seagrass meadows, estimated at 7,000 hectares in 2010 (<xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>). We confirm the presence of seagrasses in the Canal del Infiernillo; however, we cannot know the species composition. Despite this, the meadows have shown remarkable stability and density, possibly due to Comcaac&#x2019;s conservation efforts. This area is part of the Comcaac (Seri) community&#x2019;s territory, known for its historical use of seagrasses, and has been under exclusive fishing concession since 1975 (<xref ref-type="bibr" rid="B25">DOF, 1975</xref>; <xref ref-type="bibr" rid="B37">Felger et&#xa0;al., 1980</xref>; <xref ref-type="bibr" rid="B36">Felger and Moser, 1985</xref>).</p>
<p>Bah&#xed;a Concepci&#xf3;n hosts the only population of <italic>Z. marina</italic> on the east coast of the Gulf of California. Documented on two beaches, El Requeson and Punta Arenas, these meadows cover roughly 33.74 hectares (<xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>). However, during our field work in January 2023, no <italic>Zostera marina</italic> was found, only <italic>Ruppia maritima</italic> was present. This finding may be due to the increase in temperatures in the region (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>), due to <italic>Z. marina</italic> declines at temperatures above 25&#xb0;C and disappears at 28&#xb0;C, while <italic>R. maritima</italic> has a wide range of tolerance to temperature changes (<xref ref-type="bibr" rid="B89">Phillips and Me&#xf1;ez, 1988</xref>; <xref ref-type="bibr" rid="B55">Lazar and Dawes, 1991</xref>; <xref ref-type="bibr" rid="B68">Meling-Lopez and Ibarra-Obando, 1999</xref>; <xref ref-type="bibr" rid="B99">Santamar&#xed;a-Gallegos et&#xa0;al., 2000</xref>). The ecological succession between both species had already been previously reported at the this and another sites, mainly in areas of lower environmental quality (<xref ref-type="bibr" rid="B68">Meling-Lopez and Ibarra-Obando, 1999</xref>; <xref ref-type="bibr" rid="B15">Cho et al., 2009</xref>; <xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>).</p>
<p>The status of seagrass meadows on the eastern Gulf of California coast is uncertain. We can confirm the presence of seagrasses in 13 sites. However, we do not know the composition of the seagrass species, since the key meadows are found predominantly within coastal lagoons, with high levels of eutrophication. For example, Bah&#xed;a de Kino, where significant anthropogenic activities have been recorded, it is too difficult identified species using remote technologies.</p>
<p>Regarding the several new locations for <italic>H. wrightii</italic> and <italic>H. decipiens</italic> in La Paz (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 1</bold>
</xref>), we think that changes in seagrass distribution can be attributed to the reproductive strategies and seasonal dynamics, linked to life cycles, which can result in meadows disappearing for years at a time, especially among annual species (<xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>). <italic>H. wrightii</italic>, a perennial species, primarily propagates vegetatively (<xref ref-type="bibr" rid="B96">Santamar&#xed;a-Gallegos, 2016</xref>; <xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al, 2021</xref>). While <italic>H. decipiens</italic>, an annual species, germinates from seeds and is visible from April to October, declining in winter in most locations (<xref ref-type="bibr" rid="B20">Dawes and Lawrence, 1980</xref>; <xref ref-type="bibr" rid="B19">Dawes et&#xa0;al., 1995</xref>; <xref ref-type="bibr" rid="B53">Kuo and Kirkman, 1995</xref>; <xref ref-type="bibr" rid="B51">Kenworthy, 1999</xref>; <xref ref-type="bibr" rid="B23">Deis, 2000</xref>; <xref ref-type="bibr" rid="B96">Santamar&#xed;a-Gallegos, 2016</xref>). These species form transient, scattered patches that are often underreported and difficult to detect due to algal overgrowth. Misidentification with macroalgae or other seagrasses is common, which may have contributed to the overestimation of <italic>H. decipiens</italic> in the Gulf of California&#x2019;s marine flora records.</p>
<p>While documented changes in <italic>Z. marina</italic> meadows exist, other species lack comprehensive distribution data. For example<italic>, R. maritima</italic> was last officially reported in Los Cabos in the late 1990s (<xref ref-type="bibr" rid="B90">Ram&#xed;rez-Garc&#xed;a and Lot, 1994</xref>; <xref ref-type="bibr" rid="B56">Le&#xf3;n de la Luz et&#xa0;al., 1997</xref>), with no subsequent sightings. Consequently, seagrass mapping and monitoring efforts should account for their dynamic nature. Regular long-term monitoring, as suggested by <xref ref-type="bibr" rid="B112">Valle et&#xa0;al. (2013)</xref>, including both remote sensing and fieldwork over periods such as four years, could yield more accurate spatial distribution models for these populations (<xref ref-type="bibr" rid="B9">Bremner et&#xa0;al., 2023</xref>). It is crucial to conduct detailed fieldwork to revalidate species distributions and understand the ecological dynamics of seagrass meadows in the Gulf of California.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Summary of human and climate stressors</title>
<p>Coastal lagoons and estuaries along the Sonora and Sinaloa coastlines are subject to three major human-induced stressors: pollution, coastal modification, and overuse of biological resources <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>. The environmental stress in these regions is largely attributed to population growth and the indirect effects of agricultural and livestock runoff (<xref ref-type="bibr" rid="B69">Montes et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B79">P&#xe1;ez-Osuna et&#xa0;al., 2017</xref>).</p>
<p>The primary environmental threat comes from agricultural pollutants like fertilizers and pesticides. These substances accumulate in sediments and water, posing risks to the health of ecosystems and leading to significant marine pollution (<xref ref-type="bibr" rid="B14">Carvalho et al., 1996</xref>; <xref ref-type="bibr" rid="B43">Galindo-Reyes et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B79">P&#xe1;ez-Osuna et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B93">Ru&#xed;z-Fern&#xe1;ndez et&#xa0;al., 2002</xref>). In Sonora and Sinaloa, the expansion of aquaculture has also been problematic. Related coastal alterations have substantially affected local hydrology and altered coastal processes. Organic discharges and nutrient outflow from aquaculture operations diminish water quality, which in turn adversely affects vegetation cover (<xref ref-type="bibr" rid="B2">Alonso-P&#xe9;rez et&#xa0;al., 2003</xref>; <xref ref-type="bibr" rid="B46">Gonz&#xe1;lez-Abraham et&#xa0;al., 2015</xref>), and may lead to succession events in vegetation, such as those observed between <italic>R. maritima</italic> and <italic>Z. marina</italic> (<xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>).</p>
<p>In Baja California Peninsula, coastal alterations and tourism are the primary threats to seagrass meadows (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>). In Los Cabos, tourism infrastructure has led to significant ecological strain, with the San Jos&#xe9; estuary&#x2019;s geomorphology reduced by over 40% in 20 years, affecting its ecosystem services (<xref ref-type="bibr" rid="B4">Arizpe et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B117">Wurl, 2019</xref>). While, tourism development, combined with natural events like hurricanes and climate phenomena, has been detrimental, particularly to <italic>H. wrightii</italic> in La Paz Bay&#x2019;s Balandra MPA (<xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>).</p>
<p>Environmental stressors, such as pronounced warming and heatwave events, are higher in Loreto and Concepcion Bay (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>). The seagrass meadows of Bah&#xed;a Concepci&#xf3;n face a challenging environment due to their small size, human threats, nutrient discharges leading to eutrophication, and climate change impacts. Studies indicate a significant decrease in seed banks from 1995 to 2010 (<xref ref-type="bibr" rid="B99">Santamar&#xed;a-Gallegos et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>).</p>
<p>The Midriff Islands Region, including Bah&#xed;a de los &#xc1;ngeles, Tiburon, and &#xc1;ngel de la Guarda islands, faces less environmental pressure, making these sites potential climate refugia (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>). The cooler temperatures and upwelling events here present a research opportunity for understanding future climate impacts on seagrasses (<xref ref-type="bibr" rid="B57">L&#xf3;pez et&#xa0;al., 2006</xref>).</p>
<p>Despite some seagrasses being located within protected areas, their conservation is not guaranteed due to the highly disturbed and fragmented coastal landscape. As shown in <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>, few seagrass records fall within fully protected zones with restrictions on extractive activities. Most are in minimally protected or unprotected areas. If current pressures like habitat degradation and population reduction continue, some seagrass populations may risk extinction in the short to medium term. Immediate and proactive conservation measures are critical for seagrass recovery and preservation.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Conservation and management</title>
<p>Despite increased recognition of the importance of seagrass conservation, often linked with the protection of mangroves and reefs, a significant gap persists between scientific understanding and its integration into effective policy and conservation action (<xref ref-type="bibr" rid="B28">Duarte et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B60">L&#xf3;pez-Calder&#xf3;n et al., 2014</xref>; <xref ref-type="bibr" rid="B41">Fortes et&#xa0;al., 2018</xref>). For example, despite its recognition as a Priority Marine Area, granted in 1998 by CONABIO, USAID, WWF, FMCN and Packard Foundation (<xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>), Bah&#xed;a Concepci&#xf3;n lacks formal protection and a dedicated conservation strategy.</p>
<p>The complex interaction among resource management, public policy, and economic objectives in coastal regions creates a barrier to implementing laws that protect seagrass ecosystems. Developing management plans that safeguard key ecosystems while accommodating local economic interests is a delicate balance, often met with opposition to restrictions (<xref ref-type="bibr" rid="B7">Bennett and Dearden, 2014</xref>). Standard conservation strategies may not always be suitable for seagrass habitats due to their unique dynamics and fragility. For instance, the establishment of the Balandra Flora and Fauna Protection Area in November 2012 (<xref ref-type="bibr" rid="B26">DOF, 2012</xref>), which aimed to limit certain activities, paradoxically intensified tourism pressure, thereby potentially impacting the local <italic>H. wrightii</italic> population (<xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>). This development highlights the insufficient consideration given to seagrass meadows within management plans. Consequently, despite numerous seagrass locations falling within designated protection areas (refer to <xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>), there is no assurance that these habitats are effectively safeguarded.</p>
<p>Community engagement is crucial in conserving seagrass ecosystems, which are frequently undervalued and misunderstood. Successful conservation models have leveraged environmental education and citizen science to involve coastal communities in monitoring seagrass populations (<xref ref-type="bibr" rid="B18">Cunha et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B118">Yaakub et&#xa0;al., 2014</xref>). Identifying and safeguarding potential threat zones, creating micro-protected areas, and conducting water quality assessments could provide vital sanctuaries for seagrass recovery and scientific study. A multidisciplinary approach is essential for seagrass conservation, one that addresses the ecological, societal, and economic facets of these critical habitats.</p>
</sec>
<sec id="s4_4">
<label>4.4</label>
<title>Current information and knowledge gaps</title>
<p>Since 1980, the number of peer-reviewed publications and the increase in theses and reports indicate a resurgence of interest in seagrass research, led mainly by local researchers (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4A</bold>
</xref>). However, there are still notable gaps in knowledge.</p>
<p>Seagrasses have been documented since 1645, documenting the cultural and ethnobotanical uses of these plants (<xref ref-type="bibr" rid="B35">Felger and Moser, 1973</xref>; <xref ref-type="bibr" rid="B33">Felger, 1976</xref>; <xref ref-type="bibr" rid="B34">Felger and Lowe, 1976</xref>; <xref ref-type="bibr" rid="B102">Sheridan and Felger, 1977</xref>; <xref ref-type="bibr" rid="B111">Valencia et al., 1985</xref>; <xref ref-type="bibr" rid="B38">Felger et al., 2013</xref>). This type of research has been published as ethnobotanical studies and floristic lists that also provide information on distribution (<xref ref-type="bibr" rid="B24">Den Hartog, 1970</xref>; <xref ref-type="bibr" rid="B67">McMillan and Phillips, 1979b</xref>; <xref ref-type="bibr" rid="B116">Wilder et al., 2007</xref>). On the other hand, in the 1960s emerged biological and ecological studies (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>). The first topic describes mainly phenological and reproductive aspects of seagrasses (<xref ref-type="bibr" rid="B21">Dawson, 1944</xref>; <xref ref-type="bibr" rid="B101">Setchell, 1946</xref>; <xref ref-type="bibr" rid="B22">Dawson, 1960</xref>; <xref ref-type="bibr" rid="B30">Edwards, 1978</xref>; <xref ref-type="bibr" rid="B66">McMillan and Phillips, 1979a</xref>; <xref ref-type="bibr" rid="B65">McMillan, 1983</xref>; <xref ref-type="bibr" rid="B86">Phillips and Backman, 1983</xref>; <xref ref-type="bibr" rid="B87">Phillips et&#xa0;al., 1983</xref>).</p>
<p>Meanwhile the ecological issues explore interactions with other species, including algae, fishes, invertebrates (<xref ref-type="bibr" rid="B88">Phillips and McRoy, 1980</xref>; <xref ref-type="bibr" rid="B81">Paul &amp; Bowers, 1982</xref>; <xref ref-type="bibr" rid="B73">Oliva-Martinez and Ortega, 1983</xref>; <xref ref-type="bibr" rid="B85">Phillips, 1984</xref>; <xref ref-type="bibr" rid="B77">Ortega et&#xa0;al., 1986</xref>; <xref ref-type="bibr" rid="B31">Ezcurra et&#xa0;al., 1988</xref>; <xref ref-type="bibr" rid="B40">Flores-Verdugo et&#xa0;al., 1988</xref>; <xref ref-type="bibr" rid="B50">Ibarra-Obando and R&#xed;os, 1993</xref>). Other aspects addressed in ecological studies include biomass quantification, population changes, and energy flow models (<xref ref-type="bibr" rid="B95">S&#xe1;nchez-Lizaso and Riosmena Rodr&#xed;guez, 1997</xref>; <xref ref-type="bibr" rid="B106">Torre-Cosio, 2002</xref>; <xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>, <xref ref-type="bibr" rid="B60">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B113">Van Dam et&#xa0;al., 2021</xref>).</p>
<p>New critical research themes, as conservation and genetics, emerged from the 1990s, providing assessment of biodiversity threats and conservation status of the ecosystem (<xref ref-type="bibr" rid="B45">Glenn et&#xa0;al., 1992</xref>; <xref ref-type="bibr" rid="B12">Brusca et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B6">Basurto, 2008</xref>; <xref ref-type="bibr" rid="B70">Morzaira-Luna and Daneman, 2018</xref>; <xref ref-type="bibr" rid="B91">Riosmena-Rodr&#xed;guez et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>). Including the evaluation of variability, resilience, and connectivity of <italic>Z. marina</italic> populations on the Gulf of California and the Pacific coast (<xref ref-type="bibr" rid="B5">Backman, 1991</xref>; <xref ref-type="bibr" rid="B71">Mu&#xf1;&#xed;z-Salazar et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B105">Talbot et&#xa0;al., 2016</xref>) (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4B</bold>
</xref>).</p>
<p>Despite this broad spectrum of research, there exist certain areas of study that remain under-researched, representing significant knowledge gaps. For instance, seagrass distribution has been examined since the initial records on the coasts of the Gulf of California (<xref ref-type="bibr" rid="B24">Den Hartog, 1970</xref>; <xref ref-type="bibr" rid="B66">McMillan and Phillips, 1979a</xref>; <xref ref-type="bibr" rid="B1">Aguilar-Rosas and L&#xf3;pez-Ruelas, 1985</xref>; <xref ref-type="bibr" rid="B50">Ibarra-Obando and R&#xed;os, 1993</xref>; <xref ref-type="bibr" rid="B92">Riosmena-Rodr&#xed;guez and S&#xe1;nchez-Lizaso, 1996</xref>; <xref ref-type="bibr" rid="B98">Santamar&#xed;a-Gallegos et&#xa0;al., 2006</xref>). However, the depth of information varies greatly: some reports merely document species presence, while others include meadow sizes. Estimations of meadow extent are seldom reported, with the only data originating from Canal del Infiernillo, Bah&#xed;a Concepci&#xf3;n and Bah&#xed;a Balandra (<xref ref-type="bibr" rid="B106">Torre-Cosio, 2002</xref>; <xref ref-type="bibr" rid="B48">Hinojosa-Arango et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B60">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2014</xref>, <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>).</p>
<p>The subject of blue carbon in seagrass, though well-studied globally (<xref ref-type="bibr" rid="B54">Lavery et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B76">Oreska et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B42">Friess, 2023</xref>), is lacking in the Gulf of California, with only two studies providing information (<xref ref-type="bibr" rid="B39">Flores-Verdugo et&#xa0;al., 1995</xref>; <xref ref-type="bibr" rid="B47">Herrera-Silveira et&#xa0;al., 2018</xref>). Lastly, fishing and aquaculture research indirectly offer information on seagrass distribution (<xref ref-type="bibr" rid="B81">Paul and Bowers, 1982</xref>; <xref ref-type="bibr" rid="B97">Santamar&#xed;a-Gallegos et&#xa0;al., 1999</xref>; <xref ref-type="bibr" rid="B119">Zetina-Rej&#xf3;n, 1999</xref>; <xref ref-type="bibr" rid="B8">Bourill&#xf3;n-Moreno, 2002</xref>).</p>
<sec id="s4_4_1">
<label>4.4.1</label>
<title>Research by species</title>
<p>The species <italic>Z. marina</italic> has been the primary focus of most of these studies since its initial documentation in 1645 (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>). It has been the subject of at least 44 individual studies (<xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B91">Riosmena-Rodr&#xed;guez et&#xa0;al., 2013</xref>; <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>), and its presence has been noted in 15 distinct locations (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 2</bold>
</xref>). It is an annual species, which germinates during the fall, reaching its greatest abundance in the winter, mainly when the water temperature ranges between 17-19&#xb0;C. About 70% of the species&#x2019; records have been documented during the fall-winter season. Its decline begins with increasing temperatures, disappearing almost completely during the summer (<xref ref-type="bibr" rid="B86">Phillips and Backman, 1983</xref>; <xref ref-type="bibr" rid="B88">Phillips and McRoy, 1980</xref>; <xref ref-type="bibr" rid="B98">Santamar&#xed;a-Gallegos et&#xa0;al., 2006</xref>).</p>
<p>Data on the expanse of <italic>Z. marina</italic> meadows, however, is limited and only available for Canal del Infiernillo and Bah&#xed;a Concepci&#xf3;n (<xref ref-type="bibr" rid="B106">Torre-Cosio, 2002</xref>; <xref ref-type="bibr" rid="B61">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2016</xref>). However, in recent years, <italic>Z. marina</italic> has not been reported in Bah&#xed;a Concepci&#xf3;n, which can be considered an indicator that the species is at risk, at least in that region. In the Canal del Infiernillo, the populations remain apparently stable and without human stressors (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2</bold>
</xref>) under the management of the Seri culture. These populations maintain genetic connectivity between Bah&#xed;a Concepci&#xf3;n and may be the key to rescuing the populations of the Baja California peninsula (<xref ref-type="bibr" rid="B71">Mu&#xf1;&#xed;z-Salazar et&#xa0;al., 2005</xref>) through seagrass dispersal aided by water mass exchanges between these key Gulf locations (<xref ref-type="bibr" rid="B58">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2008</xref>).</p>
<p>
<italic>Ruppia maritima</italic>, with reports of its presence in 23 different localities (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 2</bold>
</xref>), is the most broadly distributed species within the Gulf of California. Though frequently studied within wetland contexts, <italic>R. maritima</italic> is often overlooked as a seagrass species (<xref ref-type="bibr" rid="B74">Oliveira et&#xa0;al., 1983</xref>; <xref ref-type="bibr" rid="B52">Kuo and den Hartog, 2001</xref>). It is a very resistant species that tolerates wide ranges of temperature and salinity; it is commonly present in areas of lower environmental quality (<xref ref-type="bibr" rid="B15">Cho et al., 2009</xref>; <xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>).</p>
<p>This species is mentioned in 29 studies, either directly or indirectly, however information on its distribution, abundance, or biomass estimation is provided in a mere six studies (<xref ref-type="bibr" rid="B30">Edwards, 1978</xref>; <xref ref-type="bibr" rid="B73">Oliva-Martinez and Ortega, 1983</xref>; <xref ref-type="bibr" rid="B40">Flores-Verdugo et&#xa0;al., 1988</xref>; <xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>, <xref ref-type="bibr" rid="B60">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B44">Garc&#xed;a-Trasvi&#xf1;a, 2017</xref>). Its rising prominence in recent publications due to increasing biomass, spatial distribution, and significance as a food source for green turtles, is noteworthy (<xref ref-type="bibr" rid="B59">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2010</xref>, <xref ref-type="bibr" rid="B60">L&#xf3;pez-Calder&#xf3;n et&#xa0;al., 2014</xref>).</p>
<p>In this study we recorded <italic>R. maritima</italic> in four new locations in the municipality of Loreto and one more in Muleg&#xe9; (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 1</bold>
</xref>). In El Juncalito and Puerto Escondido beaches the plants were short (less than 10 cm in height), lacked reproductive structures and showed low density. Green algae of the genus <italic>Caulerpa</italic> were mixed with <italic>R. maritima</italic>. Respect to the records of the estuaries, the <italic>R. maritima</italic> populations were associated to other kinds of vegetation (marshes and mangroves in the estuaries of Loreto and palm trees in Muleg&#xe9;). All of them presented conditions of isolation, however, they are threatened by human activities. This species remains the only seagrass species in Mexico that is not included in NOM-059.</p>
<p>The species <italic>H. wrightii</italic> was initially documented in the Canal del Infiernillo in 1979 and has since been reported in seven localities (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 1</bold>
</xref>). Eight studies focused on the species are reported. Preliminary studies concentrated on the presence of this species within the Gulf of California (<xref ref-type="bibr" rid="B67">McMillan and Phillips, 1979b</xref>; <xref ref-type="bibr" rid="B1">Aguilar-Rosas and L&#xf3;pez-Ruelas, 1985</xref>; <xref ref-type="bibr" rid="B90">Ram&#xed;rez-Garc&#xed;a and Lot, 1994</xref>). However, since 2010, new studies have emerged that focus on the seasonal characteristics of <italic>H. wrightii</italic>, including evaluations of the abundance and morphology of its shoots, as well as biomass quantification (<xref ref-type="bibr" rid="B96">Santamar&#xed;a-Gallegos, 2016</xref>; <xref ref-type="bibr" rid="B44">Garc&#xed;a-Trasvi&#xf1;a, 2017</xref>). Notably, <xref ref-type="bibr" rid="B13">Campos-D&#xe1;vila et&#xa0;al. (2019)</xref> and <xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al. (2021</xref>, <xref ref-type="bibr" rid="B83">2023)</xref> have provided descriptions of the species&#x2019; phenology, physiological traits, and fauna associated with <italic>H. wrightii</italic> meadows.</p>
<p>We contribute with new information to the knowledge of the species, reporting new sites within Bah&#xed;a de La Paz where it had not been reported before (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 1</bold>
</xref>). These new records, indicates an expansion of its distribution range, probably favored by its tolerance to changes in temperature and salinity (<xref ref-type="bibr" rid="B84">Phillips, 1960</xref>; <xref ref-type="bibr" rid="B63">Mazzotti et&#xa0;al., 2007</xref>). Despite this new information, a decrease in the extent of the patches has also been observed. For example, in Balandra Bay, a reduction of 70% is estimated between 2013 and the present (<xref ref-type="bibr" rid="B82">P&#xe9;rez-Estrada et&#xa0;al., 2021</xref>).</p>
<p>Finally, <italic>H. decipiens</italic> has been documented exclusively in Bah&#xed;a de La Paz, including in Ensenada de La Paz and in estuaries on the West side of Isla Esp&#xed;rito Santo (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Datasheet 3</bold>
</xref>). The three existing studies on this species have described the seasonal variation in its life cycle, its characterization, and biomass quantification (<xref ref-type="bibr" rid="B98">Santamar&#xed;a-Gallegos et&#xa0;al., 2006</xref>; <xref ref-type="bibr" rid="B96">Santamar&#xed;a-Gallegos, 2016</xref>; <xref ref-type="bibr" rid="B44">Garc&#xed;a-Trasvi&#xf1;a, 2017</xref>). As far as we know, it remains a species restricted to Bah&#xed;a de La Paz. However, although in the past it had been reported exclusively on Costa Baja beach (<xref ref-type="bibr" rid="B98">Santamar&#xed;a-Gallegos et&#xa0;al., 2006</xref>), this work adds new records (<xref ref-type="supplementary-material" rid="SM1">
<bold>Supplementary Table 1</bold>
</xref>). Indicating, as in the case of <italic>H. wrightii</italic>, an expansion in its distribution range, probably driven by its tropical affinity (<xref ref-type="bibr" rid="B98">Santamar&#xed;a-Gallegos et&#xa0;al., 2006</xref>).</p>
</sec>
</sec>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>This review represents an initial comprehensive assessment of the conservation status and knowledge of seagrass species in the Gulf of California, pointing to a significant lag in research in the region and the need to incentivize investment by government institutions. To address knowledge gaps and advance conservation, the following measures are recommended:</p>
<list list-type="bullet">
<list-item>
<p>Formation of a collaborative network among seagrass ecosystem researchers to foster data sharing and joint studies.</p>
</list-item>
<list-item>
<p>Regular updates on mapping the distribution and area of seagrass species, particularly within MPAs, utilizing a combination of remote sensing and fieldwork, including area estimation and use of quadrats for sampling. The differentiation between seagrass patches and meadows must be clear and consistent.</p>
</list-item>
<list-item>
<p>Implementation of environmental programs to improve assessment of ecosystem services, with a focus on key environmental indicators like temperature, water quality, nutrient levels, and heavy metal contents to gauge ecosystem health.</p>
</list-item>
<list-item>
<p>Detailed investigation into the phenological and reproductive patterns of seagrass species in the Gulf to comprehend their transient nature, complemented by genetic research to understand population connectivity and gene flow.</p>
</list-item>
<list-item>
<p>Assessment of seagrass contributions to fisheries, including studies on fauna-associated dispersal and the effects of atmospheric phenomena on these ecosystems.</p>
</list-item>
<list-item>
<p>In-depth exploration of biogeochemical cycling, carbon storage, and sequestration within seagrass habitats to understand their role in climate mitigation.</p>
</list-item>
<list-item>
<p>Implementation of environmental educations program that raises awareness among the population about the importance of seagrasses. Forums and workshops can be tools that activate participation through citizen science.</p>
</list-item>
</list>
</sec>
<sec id="s6" sec-type="author-contributions">
<title>Author contributions</title>
<p>MR-Z: Conceptualization, Data curation, Formal analysis, Investigation, Methodology, Software, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing. CP-E: Investigation, Methodology, Resources, Validation, Writing &#x2013; review &amp; editing. JL-C: Data curation, Investigation, Resources, Writing &#x2013; review &amp; editing. AC: Data curation, Resources, Writing &#x2013; review &amp; editing. SV: Data curation, Resources, Validation, Writing &#x2013; review &amp; editing. FF: Conceptualization, Funding acquisition, Investigation, Project administration, Resources, Supervision, Validation, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing.</p>
</sec>
</body>
<back>
<sec id="s7" sec-type="funding-information">
<title>Funding</title>
<p>The author(s) declare that no financial support was received for the research, authorship, and/or publication of this article.</p>
</sec>
<ack>
<title>Acknowledgments</title>
<p>MRZ and FF thank the support of the Institute of the Americas, in particular, Richard Kiy, Tania Miranda and Catalina L&#xf3;pez Sagastegui.</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="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="s10" 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.2024.1402044/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2024.1402044/full#supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="Table_1.pdf" id="ST1" mimetype="application/pdf"/>
<supplementary-material xlink:href="Table_2.pdf" id="ST2" mimetype="application/pdf"/>
<supplementary-material xlink:href="DataSheet_1.xlsx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
<supplementary-material xlink:href="DataSheet_2.pdf" id="SM2" mimetype="application/pdf"/>
<supplementary-material xlink:href="DataSheet_3.xlsx" id="SM3" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"/>
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