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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.2024.1466834</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>Coral reefs of Pakistan: a comprehensive review of anthropogenic threats, climate change, and conservation status</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name>
<surname>Ahmad</surname>
<given-names>Ishfaq</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
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<contrib contrib-type="author">
<name>
<surname>Guo</surname>
<given-names>Pu</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="author-notes" rid="fn003">
<sup>&#x2020;</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Zhao</surname>
<given-names>Mei-Xia</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff3">
<sup>3</sup>
</xref>
<xref ref-type="author-notes" rid="fn001">
<sup>*</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/1046433"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhong</surname>
<given-names>Yu</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Zheng</surname>
<given-names>Xiao-Yun</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Zhang</surname>
<given-names>Shu-Qi</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Qiu</surname>
<given-names>Jian-Wen</given-names>
</name>
<xref ref-type="aff" rid="aff4">
<sup>4</sup>
</xref>
<xref ref-type="aff" rid="aff5">
<sup>5</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/131192"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Shi</surname>
<given-names>Qi</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/928076"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Yan</surname>
<given-names>Hong-Qiang</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2572352"/>
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</contrib>
<contrib contrib-type="author">
<name>
<surname>Tao</surname>
<given-names>Shi-Chen</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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<contrib contrib-type="author">
<name>
<surname>Xu</surname>
<given-names>Li-Jia</given-names>
</name>
<xref ref-type="aff" rid="aff6">
<sup>6</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/writing-review-editing/"/>
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</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Key Laboratory of Ocean and Marginal Sea Geology, South China Sea Institute of Oceanology, Chinese Academy of Sciences</institution>, <addr-line>Guangzhou</addr-line>, <country>China</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>College of Marine Sciences, University of Chinese Academy of Sciences</institution>, <addr-line>Beijing</addr-line>, <country>China</country>
</aff>
<aff id="aff3">
<sup>3</sup>
<institution>State Key Laboratory of Marine Environmental Science, Xiamen University</institution>, <addr-line>Xiamen</addr-line>, <country>China</country>
</aff>
<aff id="aff4">
<sup>4</sup>
<institution>Department of Biology, Hong Kong Baptist University</institution>, <addr-line>Hong Kong</addr-line>, <country>China</country>
</aff>
<aff id="aff5">
<sup>5</sup>
<institution>Hong Kong Branch of the Southern Marine Science and Engineering Guangdong Laboratory (Guangzhou), Hong Kong University of Science and Technology</institution>, <addr-line>Hong Kong</addr-line>, <country>China</country>
</aff>
<aff id="aff6">
<sup>6</sup>
<institution>Research Centre for Marine Ecology and Environment, South China Institute of Environmental Sciences</institution>, <addr-line>Guangzhou</addr-line>, <country>China</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>Edited by: Robert Mortimer, York St. John University, United Kingdom</p>
</fn>
<fn fn-type="edited-by">
<p>Reviewed by: Fabio Favoretto, University of California, San Diego, United States</p>
<p>Abhra Chanda, Jadavpur University, India</p>
</fn>
<fn fn-type="corresp" id="fn001">
<p>*Correspondence: Mei-Xia Zhao, <email xlink:href="mailto:zhaomeix@scsio.ac.cn">zhaomeix@scsio.ac.cn</email>
</p>
</fn>
<fn fn-type="other" id="fn003">
<p>&#x2020;These authors share first authorship</p>
</fn>
</author-notes>
<pub-date pub-type="epub">
<day>30</day>
<month>09</month>
<year>2024</year>
</pub-date>
<pub-date pub-type="collection">
<year>2024</year>
</pub-date>
<volume>11</volume>
<elocation-id>1466834</elocation-id>
<history>
<date date-type="received">
<day>18</day>
<month>07</month>
<year>2024</year>
</date>
<date date-type="accepted">
<day>04</day>
<month>09</month>
<year>2024</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2024 Ahmad, Guo, Zhao, Zhong, Zheng, Zhang, Qiu, Shi, Yan, Tao and Xu</copyright-statement>
<copyright-year>2024</copyright-year>
<copyright-holder>Ahmad, Guo, Zhao, Zhong, Zheng, Zhang, Qiu, Shi, Yan, Tao and Xu</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>The coral reefs along Pakistan&#x2019;s coastline are of ecological and economic significance. However, they are increasingly threatened by anthropogenic threats and climate change. These vulnerable ecosystems are declining due to various factors such as sedimentation, climate change, overfishing, destructive fishing practices, marine pollution, and tourism development. We found that 29%, 24%, 26%, 16% and 18% of the studies exceeding, marine pollution, overfishing/destructive fishing, coastal tourism, climate change and sedimentation, respectively; thus, indicating inadequate water quality status in part of Pakistan coastal water. These influences lead to several negative impacts, such as jeopardized coral health, decline in biodiversity, and the simplification of reef structures. In response to these threats, conservation efforts are imperative. This literature review provides an in-depth analysis of anthropogenic threats, climate change and the conservation of coral reefs in Pakistan. This review provides suggestions on how the country could better conserve its coral reef ecosystem. These include (1) initiatives such as establishing marine protected areas (MPAs), (2) encouraging sustainable fishing practices and reducing pollution, (3) developing the country as an ecotourism destination and implementing climate change adaptation measures, and (4) community engagement through awareness campaigns and fostering collaboration among, governmental organizations, non-governmental organizations (NGOs) and scientists. These comprehensive conservation policies address human-caused and other challenges, safeguarding their ecological, economic, and cultural relevance for future generations.</p>
</abstract>
<kwd-group>
<kwd>marine protected areas (MPAs)</kwd>
<kwd>coral reef restoration</kwd>
<kwd>ecosystem</kwd>
<kwd>marine pollution</kwd>
<kwd>overfishing</kwd>
<kwd>sedimentation</kwd>
</kwd-group>
<counts>
<fig-count count="6"/>
<table-count count="1"/>
<equation-count count="0"/>
<ref-count count="153"/>
<page-count count="16"/>
<word-count count="7099"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-in-acceptance</meta-name>
<meta-value>Coastal Ocean Processes</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>Coral reefs epitomize one of the most aesthetically captivating, ancient, and complex ecosystems on earth (<xref ref-type="bibr" rid="B86">Majumdar et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B111">Oakley-Cogan et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B112">Oppermann, 2024</xref>). They support the highest marine biodiversity, although they cover only approximately 2284,300 km<sup>2</sup>, which accounts for merely 0.09% of the total surface area of the world&#x2019;s oceans (<xref ref-type="bibr" rid="B86">Majumdar et&#xa0;al., 2018</xref>). Coral reefs provide a variety of ecological functions and services, and play a significant role in contributing billions of dollars per year through different activities such as coastal protection, tourism, food, and other goods (<xref ref-type="bibr" rid="B139">Sully et&#xa0;al., 2022</xref>). However, coral reefs are fragile ecosystems and most of them have faced serious threats in the decades (<xref ref-type="bibr" rid="B35">Burke et&#xa0;al., 2011</xref>; <xref ref-type="bibr" rid="B25">Bayraktarov et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B137">Smallhorn-West et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B46">Devlin, 2022</xref>). Recent statistics show that since the 1950s, the ability of coral reefs to provide ecosystem services has decreased by 50% (<xref ref-type="bibr" rid="B51">Eddy et&#xa0;al., 2021</xref>). Such a massive decline was caused by mounting environmental pressures from both global-scale effects and local-level impacts, which include elevated sea surface temperature (SST), sea level rise, and increased anthropogenic disruptions such as marine pollution, overfishing and destructive fishing, coastal tourism, and terrigenous sediments (<xref ref-type="bibr" rid="B126">Roberts, 1995</xref>; <xref ref-type="bibr" rid="B88">Mcclanahan, 2002</xref>; <xref ref-type="bibr" rid="B32">Brainard, 2008</xref>; <xref ref-type="bibr" rid="B45">Clark, 2009</xref>; <xref ref-type="bibr" rid="B149">Wenger et&#xa0;al., 2015</xref>). Assessing the conditions of coral reef ecology and conservation, as well as identifying threats, would provide significant support for the protection and management of these valuable marine resources.</p>
<p>In Pakistan, <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref> underscored a concerning trend of degradation in coral reefs over recent years. These reefs are mainly found along the 900 km long coastal area of Sindh and Balochistan, making it an important coastal belt, and fostering diverse ecosystems (<xref ref-type="bibr" rid="B16">Amjad and Rizvi, 1999</xref>; <xref ref-type="bibr" rid="B66">Iqbal et&#xa0;al., 2018</xref>). Coral reef communities are distributed at (a) Churna Island, near Karachi; (b) Mubarak Village; (c) Hawks Bay; (d) French Beach; (e) Sandspit; (f) Buleji; (g) Goth Abdul Rehman; (h) Ormara Rodrigues Shoals; (i) Astola Island; and (j) Jiwani (<xref ref-type="bibr" rid="B11">Ali et&#xa0;al., 2014</xref>) (<xref ref-type="fig" rid="f1">
<bold>Figure&#xa0;1</bold>
</xref>). The high biodiversity of coral reefs in Pakistan has been revealed by the gradual deepening of investigation and research in the last two decades. For example, <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref> discovered 50 live coral species in Pakistan&#x2019;s coastal waters, among which Churna Island has the highest diversity. The abovementioned localities have a strong connection with human settlements, and these coral reefs are under significant harm due to human activities. Approximately 10% of Pakistan&#x2019;s population lives in coastal areas, with 20% of these regions being somewhat developed, while about 40% of the industry is in these zones (<xref ref-type="bibr" rid="B109">Noor, 2023</xref>). Therefore major factors contributing to coral reef loss include the establishment of industrial areas, pollution, overfishing, the effects of tourism, increased sedimentation, and climate change (<xref ref-type="bibr" rid="B127">Saher et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B13">Ali et&#xa0;al., 2020</xref>, <xref ref-type="bibr" rid="B14">2021</xref>; <xref ref-type="bibr" rid="B110">Noor et&#xa0;al., 2023</xref>). Several coral conservation and restoration initiatives are currently operating in Pakistan and are overseen by various organizations, regional communities, and local and national government bodies (e.g., the Marine Fisheries Department, National Institute of Oceanography (NIO), International Union for Conservation of Nature (IUCN) (Ahmed <xref ref-type="bibr" rid="B75">Khan, 2011</xref>; <xref ref-type="bibr" rid="B94">Mohsin et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). However, the evolving dynamics of coral reef assemblages, which are largely influenced by human threats, increased sedimentation rates, and climate change, pose significant questions for conservation scientists and natural resource managers. Given the multitude of impacts expected to affect coral reefs in Pakistan in the future, this review seeks to address the primary question &#x2018;How have anthropogenic threats and climate change impacts, along with their interactions, affected coral reefs in Pakistan?&#x2019; Another question we interrogate is &#x2018;What actions should be taken to protect coral reef resources in Pakistan&#x201d;?</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>The coral reef distribution map of Pakistan.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g001.tif"/>
</fig>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<p>The literature for this study was compiled primarily from online databases and search engines, including Google Scholar (<ext-link ext-link-type="uri" xlink:href="https://scholar.google.com/">https://scholar.google.com/</ext-link>), Web of Science (<ext-link ext-link-type="uri" xlink:href="https://www.webofknowledge.com/">https://www.webofknowledge.com/</ext-link>), PubMed (<ext-link ext-link-type="uri" xlink:href="https://pubmed.ncbi.nlm.nih.gov/">https://pubmed.ncbi.nlm.nih.gov/</ext-link>) and Science Direct (sciencedirect.com). Keywords such as &#x201c;coral reefs,&#x201d; &#x201c;Pakistan,&#x201d; &#x201c;anthropogenic threats,&#x201d; &#x201c;climate change,&#x201d; &#x201c;ocean acidification,&#x201d; &#x201c;overfishing,&#x201d; &#x201c;coastal tourism,&#x201d; eutrophication,&#x201d; &#x201c;marine pollution,&#x201d; &#x201c;coral bleaching,&#x201d; &#x201c;MPAs,&#x201d; &#x201c;restoration,&#x201d; &#x201c;conservation,&#x201d; &#x201c;estuaries,&#x201d; and &#x201c;coastal water ecosystem&#x201d; were used for the search. Relevant journals in the fields of geology, marine geology, biology, marine biology, and environmental sciences were also consulted. Results from these libraries were imported to R (<xref ref-type="bibr" rid="B125">R Core Team, 2022</xref>) and subjected to analysis using the Bibliometrix package (<xref ref-type="bibr" rid="B19">Aria and Cuccurullo, 2017</xref>). A total of 136 scientific articles published between 1997 and 2023 were collected for our analysis and discussion on the coral reef ecosystems in Pakistan. The trend depicted in (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2A</bold>
</xref>) indicates a consistent annual increase in the number of published articles every year. Based on specific anthropogenic threats we describe in this paper, most of the research focuses on marine pollution as their primary parameter to measure (29%), followed by overfishing (24%), coastal tourism (26%), sedimentation (18%), climate change (16%) and other related papers (23%) (<xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2B</bold>
</xref>). Additionally, we found that specific author keywords tended to occur more frequently during a certain period. <xref ref-type="fig" rid="f2">
<bold>Figure&#xa0;2C</bold>
</xref> shows the yearly occurrences of the main terms from 1997 to 2023, but some of them increased more dynamically compared than others. The terms with the greatest increase in use over time were &#x2018;coral reefs&#x2019;, &#x2018;coral bleaching&#x2019;, &#x2018;marine pollution&#x2019;, &#x2018;MPAs&#x2019;, &#x2018;overfishing&#x2019;, &#x2018;coastal tourism&#x2019;, &#x2018;climate change&#x2019;, &#x2018;Pakistan&#x2019;, &#x2018;conservation&#x2019;, and &#x2018;restoration. Also, an extensive review of international, federal and provincial policy documents was carried out to identify those relevant to coral reef or coastal ecosystem conservation or restoration (see <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>).</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>
<bold>(A)</bold> Annual scientific production over the period from 1997 to 2023, <bold>(B)</bold> The number of articles published over time, <bold>(C)</bold> Analysis of the top 10 keywords from 1997 to 2023.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g002.tif"/>
</fig>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Initiatives for coral reef conservation in Pakistan.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="top" align="left">Institution/initiative</th>
<th valign="top" align="left">Description</th>
<th valign="top" align="left">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="top" align="left">
<bold>National Institute of Oceanography (NIO)</bold>
</td>
<td valign="top" align="left">This department performing research and studies on marine and coastal ecosystems plays an important role in monitoring and conserving coral reefs and other marine ecosystems.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B115">Pakistan, 2007</xref>; <xref ref-type="bibr" rid="B75">Khan, 2011</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">
<bold>National Centre for Maritime Policy Research (NCMPR)</bold>
</td>
<td valign="top" align="left">The National Center for Maritime Policy Research focuses on conducting research related to maritime issues, with a primary emphasis on marine conservation.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B4">Ahmad, 2015</xref>; <xref ref-type="bibr" rid="B23">Azmie, 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">
<bold>International Union for Conservation of Nature (IUCN) Pakistan</bold>
</td>
<td valign="top" align="left">IUCN runs several initiatives related to the conservation of maritime ecosystems. They collaborate with stakeholders and local people to promote sustainable fisheries and the protection of maritime habitats.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B92">Memon, 2012</xref>; <xref ref-type="bibr" rid="B97">Mukhtar and Hannan, 2012</xref>; <xref ref-type="bibr" rid="B123">Rafique, 2018</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">
<bold>Sindh Coastal and Inland Community Development Project (SCICDP)</bold>
</td>
<td valign="top" align="left">This project is funded by the Sindh Rural Support Program (PRSP). The project aims to enhance the livelihood of coastal communities particularly fisheries and conservation of marine biodiversity.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B141">Tambunan, 2004</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">
<bold>Balochistan Coastal Development Authority (BCDA)</bold>
</td>
<td valign="top" align="left">The BCDA initiatives focus on the conservation of coastal ecosystems in Balochistan.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B68">Jamali, 2009</xref>; <xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>)</td>
</tr>
<tr>
<td valign="top" align="left">
<bold>Marine Fisheries Department</bold>
</td>
<td valign="top" align="left">Fisheries department monitoring and regulating activities along the country&#x2019;s coastline. This department plays a significant role in the management of Pakistan&#x2019;s marine resources and the implementation of sustainable practices.</td>
<td valign="top" align="left">(<xref ref-type="bibr" rid="B94">Mohsin et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Finally, the collected literature was critically reviewed, during which significant information regarding the introduction, research status of coral reefs in Pakistan, anthropogenic threats, coral reef restoration, and conservation efforts was obtained, and the research gaps were carefully extracted and synthesized in the main sections of the review paper. The findings and insights in this review are based on the scientific knowledge available at the time of writing, and every effort has been made to ensure the accuracy of the information presented.</p>
</sec>
<sec id="s3">
<label>3</label>
<title>Ecological status of coral reefs in Pakistan</title>
<p>Coral reefs in Pakistan are distributed in the southeastern part of the Arabian Sea (<xref ref-type="bibr" rid="B150">Wilson et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B11">Ali et&#xa0;al., 2014</xref>). However, the research on corals in Pakistan is still comparatively a more recent phenomenon. In 2006, a field visit conducted by the Pakistan Wetlands Programme (PWP), led by a coral expert from Millport University, UK, identified the first-ever coral reef habitat near Astola Island, Balochistan (<xref ref-type="bibr" rid="B78">Khan, 2011</xref>). Before this discovery, it was considered that corals do not exist within Pakistan&#x2019;s jurisdiction of the Arabian Sea.</p>
<p>Furthermore, research conducted by <xref ref-type="bibr" rid="B11">Ali et&#xa0;al. (2014)</xref> revealed that twenty-nine hard coral species (eight families and fourteen genera), one black coral species, and eight soft coral species (three families and seven genera) were identified across eighteen dive sites located in nine distinct places along the coastline of Pakistan (<xref ref-type="fig" rid="f3">
<bold>Figure&#xa0;3</bold>
</xref>). Subsequent studies by <xref ref-type="bibr" rid="B12">Ali et&#xa0;al. (2017)</xref> revealed the existence of 48 fossil coral species in the uplifted ancient coral reefs along the Balochistan coast. Additionally, a study by <xref ref-type="bibr" rid="B14">Ali et&#xa0;al. (2021)</xref> reported 58 species of fish species in the reef habitat along Churna Island. With the high biodiversity of coral communities, its ecological importance of the coral reef ecosystems has been received more and more attention (<xref ref-type="bibr" rid="B15">Amjad et&#xa0;al., 2020</xref>). However these coral reefs in Pakistan, particularly near Astola Island in Baluchistan and Churna Island in Sindh are under threats from anthropogenic threats and other natural changes (<xref ref-type="bibr" rid="B21">Aslam et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B40">Chanda, 2022</xref>). For example, according to <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref> if further shifts in community structure continue, it appears that only <italic>Porite</italic> species will survive in Pakistan&#x2019;s coastal waters. Therefore, Churna Island has been recommended for designation as an MPA because of its ecological significance (<xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Hard coral species diversity in Pakistan coastal waters [modified after <xref ref-type="bibr" rid="B11">Ali et&#xa0;al. (2014)</xref>]: <bold>(A)</bold> <italic>Goniopora albiconus</italic>; <bold>(B)</bold> <italic>Goniopora columna</italic>; <bold>(C)</bold> <italic>Goniopora djiboutiensis</italic>; <bold>(D)</bold> <italic>Goniopora albiconus</italic>; <bold>(E)</bold> <italic>Goniopora columna;</italic> <bold>(F)</bold> <italic>Goniopora djiboutiensis;</italic> <bold>(G)</bold> <italic>Goniopora cf. savignyi</italic>; <bold>(H)</bold> <italic>Goniopora somaliensis</italic>; <bold>(I)</bold> <italic>Porites harrisoni</italic>; <bold>(J)</bold> <italic>Coscinaraea</italic> sp.; <bold>(K)</bold> <italic>Psammocora obtusangulata</italic>; <bold>(L)</bold> <italic>Psammocora superficialis</italic>; <bold>(M)</bold> <italic>Porites lobata/lutea</italic>; <bold>(N)</bold> <italic>Porites monticulosa</italic>; <bold>(O)</bold> <italic>Porites nodifera</italic>; <bold>(P)</bold> <italic>Porites solida</italic>; <bold>(Q)</bold> <italic>Psammocora</italic> sp.; <bold>(R)</bold> <italic>Dendrophyllia robusta</italic>; <bold>(S)</bold> <italic>Turbinaria</italic> sp.; <bold>(T)</bold> <italic>Acanthastrea hillae</italic>; <bold>(U)</bold> <italic>Alveopora</italic> sp.; <bold>(V)</bold> <italic>Favites complanata</italic>; <bold>(W)</bold> <italic>Favites pentagona</italic>; <bold>(X)</bold> <italic>Favites spinosa</italic>; <bold>(Y)</bold> <italic>Leptastrea bottae</italic>; <bold>(Z)</bold> Acanthastrea maxima; <bold>(AA)</bold> <italic>Pocillopora damicornis</italic>; <bold>(AB)</bold> <italic>Montipora mollis</italic>; <bold>(AC)</bold> <italic>Pavona explanulata</italic>; <bold>(AD)</bold> <italic>Antipathes</italic> sp. Reprinted in a modified form with permission from Journal of the Marine Biological Association of the United Kingdom, 94(1), 75-84. <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1017/S0025315413001203">https://doi.org/10.1017/S0025315413001203</ext-link>. Copyright <sup>&#xa9;</sup> 2013 Marine Biological Association of the United Kingdom.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g003.tif"/>
</fig>
</sec>
<sec id="s4">
<label>4</label>
<title>Threats impacting coral reefs in Pakistan</title>
<p>In Pakistan, according to <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref> coral reefs have experienced the degradation in recent few years, as shown in (<xref ref-type="fig" rid="f4">
<bold>Figure&#xa0;4</bold>
</xref>), with extensive patches of bleaching in some areas in the northeastern region of Churna Island in October 2020 (<xref ref-type="bibr" rid="B67">Jamal, 2020</xref>). Therefore, anthropogenic threats and other challenges discussed below underscore the persistent difficulties confronting coral reefs in Pakistan to highlight the critical need for conservation and management efforts.</p>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Coral reef degradation in Pakistan: <bold>(A, B, E)</bold> caused by thermal stress leading to coral bleaching, and <bold>(C, D)</bold> caused by bioerosion [modified after <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref>; <xref ref-type="bibr" rid="B67">Jamal (2020)</xref>]. The red circles in the pictures clearly emphasize the areas where degradation is attributed to bioerosion.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g004.tif"/>
</fig>
<sec id="s4_1">
<label>4.1</label>
<title>Overfishing and destructive fishing</title>
<p>The coastal communities of Pakistan, particularly along the 682 miles from Sir Creek in Sindh to Jiwani in Balochistan, have relied on fishing as their primary source of food and livelihood (<xref ref-type="bibr" rid="B117">Pakistan, 2016</xref>; <xref ref-type="bibr" rid="B29">Biagi and Starnini, 2020</xref>). The marine fishing industry in Pakistan provides direct livelihoods for 125,000 people and indirectly provides nearly one million people, including dependents (<xref ref-type="bibr" rid="B77">Khan and Khan, 2011</xref>). The fish populations in the two marine provinces vary due to the differences in their topography. The majority of the fishing community is located in Karachi, Sindh, the most populous industrial city with over 18 million residents (<xref ref-type="bibr" rid="B70">Jilani, 2018</xref>), and other concentrations are in Thatta, Sindh; Pasni, Gwadar, Balochistan (<xref ref-type="bibr" rid="B1">Abbas et&#xa0;al., 2013</xref>). This coastal stretch features four harbors and nine significant landing sites and jetties (for details, see <xref ref-type="bibr" rid="B30">Birwani et&#xa0;al. (1999)</xref>. The fishing industry is supported by a fleet of more than 15,000 vessels of various sizes, including medium and small-sized boats, as well as large launches, and trawlers (<xref ref-type="bibr" rid="B77">Khan and Khan, 2011</xref>). The smaller boats, ranging from 18 to 25 feet are used for fishing in coastal waters (<xref ref-type="bibr" rid="B134">Siddiqui and Amir, 2010</xref>). Fishermen utilize hand-cast shrimp nets (known as <italic>thukri or phat</italic>) while medium-sized boats, ranging between 25 to 35 feet, use gill nets (locally known as &#x201c;<italic>rach lara&#x201d;, &#x201c;lathay ka ban&#x201d;</italic>) (<xref ref-type="bibr" rid="B77">Khan and Khan, 2011</xref>). Despite the extensive fishing activities, the fisheries sector plays a minimal contribution in Pakistan&#x2019;s economy, contributing 0.32% to the national GDP and 1.4% to the agricultural GDP, whereas employment from fisheries is minimal, at only 0.01% (<xref ref-type="bibr" rid="B148">Wasim and Abbas, 2024</xref>). In 2022, fish production reached 800,000 metric tons, only 25% was exported, generating 496 million US dollars (<xref ref-type="bibr" rid="B148">Wasim and Abbas, 2024</xref>). In response, the fishing industry in Pakistan has grown quickly, resulting in overfishing and destructive fining in several coastal and offshore areas (<xref ref-type="bibr" rid="B117">Pakistan, 2016</xref>). It is worth noting, however, that many of the existing reports are from heavily fished areas, such as coastal areas of Sindh and Balochistan (<xref ref-type="bibr" rid="B132">Siddiqi, 1992</xref>; <xref ref-type="bibr" rid="B7">Ajazuddin and Moazzam, 2007</xref>; <xref ref-type="bibr" rid="B134">Siddiqui and Amir, 2010</xref>; <xref ref-type="bibr" rid="B77">Khan and Khan, 2011</xref>; <xref ref-type="bibr" rid="B59">Gore et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B104">Nawaz and Moazzam, 2014</xref>; <xref ref-type="bibr" rid="B80">Kidwai et&#xa0;al., 2019</xref>). Karachi Fish Harbour, the country&#x2019;s primary landing area, is already overcrowded. Despite being equipped with essential facilities like a net-mending shed, chill room, and flake ice factory, the harbor faces significant challenges (<xref ref-type="bibr" rid="B80">Kidwai et&#xa0;al., 2019</xref>). The harbor&#x2019;s turning basin is large, but the area is heavily polluted. Ironically, the seawater, despite its contamination, is still used to clean the harbor floor. The handling of decaying fish contributes to making Karachi Fish Harbour one of the dirtiest in Pakistan. Despite efforts to expand the harbor&#x2019;s facilities, there is general concern that these changes will not improve the harbor&#x2019;s functionality or address the existing unsanitary conditions (<xref ref-type="bibr" rid="B104">Nawaz and Moazzam, 2014</xref>). Compounding the issue, Local fishing boats avoid using the Korangi Fish Harbour, as it was designed for vessels with drafts exceeding five meters (<xref ref-type="bibr" rid="B148">Wasim and Abbas, 2024</xref>). Similarly, Gwadar Port has been deemed unsuitable for local fishing boats. The flood of 2022 devastated most of the floating jetties in Sindh province, leaving the region without adequate landing facilities (<xref ref-type="bibr" rid="B148">Wasim and Abbas, 2024</xref>). Additionally, no landing site in the country has a functioning cold chain maintenance system. The stagnant trend in fish production, particularly in Sindh, signals that fish stocks are being severely overfished (<xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>). From 2009 to 2015, the Government of Pakistan&#x2019;s Marine Fisheries Department, in collaboration with the FAO, conducted several fish stock assessment surveys (<xref ref-type="bibr" rid="B69">Jayabalan et&#xa0;al., 2021</xref>). The final report states that large fishing fleets and current practices are &#x201c;fishing for catastrophe,&#x201d; with important fish populations in Pakistani waters decreased by 60-90 percent (<xref ref-type="bibr" rid="B148">Wasim and Abbas, 2024</xref>). This drastic decline highlights significant ecological changes and a concerning reduction in marine species. Also, according to <xref ref-type="bibr" rid="B14">Ali et&#xa0;al. (2021)</xref> these coral-associated fish communities in Pakistan are facing a threatened environment and MPAs should be prioritized in conservation efforts can greatly improve environmental and ecosystem management, giving these fish communities a great chance of surviving and thriving.</p>
</sec>
<sec id="s4_2">
<label>4.2</label>
<title>Marine pollution/contamination</title>
<p>Pollution in coastal waters is receiving attention worldwide, whereas Pakistan&#x2019;s coastal areas have been given limited attention, despite being significantly affected by anthropogenic threats and environmental challenges (<xref ref-type="bibr" rid="B127">Saher et&#xa0;al., 2019</xref>). In Pakistan, the pollution threat on the Balochistan coast is currently low because there are no major cities or large-scale projects along the coastline. Similarly, pollution levels in the delta are comparatively lower because there is no direct pollution in the area. However, the coast of Karachi is facing a significant threat from pollution generated by industry, municipal wastewater, the shipbreaking industry, and plastic waste (<xref ref-type="bibr" rid="B13">Ali et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B96">Mukheed and Alisha, 2020</xref>). Karachi, home to more than half of the country&#x2019;s industrial units and 70% of the international trade being controlled by this city, experienced a dramatic increase of population from approximately 10,000 during the 1990s and has now gone beyond 30,000 (<xref ref-type="bibr" rid="B123">Rafique, 2018</xref>). Various studies have profiled Karachi&#x2019;s coastal belt based on pollution levels (<xref ref-type="bibr" rid="B85">Majeed et&#xa0;al., 2021a</xref>, <xref ref-type="bibr" rid="B84">b</xref>; <xref ref-type="bibr" rid="B105">Neelam, 2021</xref>; <xref ref-type="bibr" rid="B107">Nergis et&#xa0;al., 2021</xref>). The accumulation of hospital waste, single-use plastics, industrial effluents and nuclear discards, contribute significantly to coastal pollution (<xref ref-type="bibr" rid="B117">Pakistan, 2016</xref>). Tanneries also release chemically rich effluents into coastal waters, with heavy metals from these sources concentrating in marine life and ultimately biomagnifying in humans, leading to toxicity (<xref ref-type="bibr" rid="B65">Iqbal and Heidegger, 2013</xref>; <xref ref-type="bibr" rid="B72">Karim, 2017</xref>). A study (<xref ref-type="bibr" rid="B84">Majeed et&#xa0;al., 2021b</xref>) assessed pollution levels at Karachi&#x2019;s ports, revealing heavy metal contamination in coastal waters from port and harbor operations. Similarly, another study (<xref ref-type="bibr" rid="B8">Alamgir et&#xa0;al., 2019</xref>) found water samples from the Lyari River mouth, Fish Harbor, and KPT boat-building area to be highly contaminated with heavy metals. The major contributors to this pollution are eutrophication and plastic waste, wreaking havoc on the fragile ecosystems along the coast of Karachi and Gaddani (<xref ref-type="bibr" rid="B153">Zuo et&#xa0;al., 2023</xref>). Eutrophication causes a significant threat to coral reefs due to excess nutrients such as nitrogen and phosphorus in the marine environment (<xref ref-type="bibr" rid="B28">Bell, 1991</xref>; <xref ref-type="bibr" rid="B50">Dorgham, 2014</xref>; <xref ref-type="bibr" rid="B103">Naumann et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B9">Albright, 2018</xref>). Algal blooms that outcompete corals for space and light, eventually smothering the corals and depriving them of essential sunlight for photosynthesis (<xref ref-type="bibr" rid="B89">Mccook, 2001</xref>; <xref ref-type="bibr" rid="B62">Hernandez-Pacheco et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B149">Wenger et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B47">Devlin and Brodie, 2023</xref>). Excessive algal growth in coral reefs not only causes ecological issues but also exacerbates plastic waste pollution, since the algae trap and accumulate plastic trash, creating a complex mess that threatens the survival of these ecosystems (<xref ref-type="bibr" rid="B60">Gregory, 2009</xref>; <xref ref-type="bibr" rid="B140">Sweet, 2019</xref>). These nutrients come from different sources such as agricultural runoff, sewage discharge, and industrial processes (<xref ref-type="bibr" rid="B17">Ansari et&#xa0;al., 2011</xref>). In Pakistan, the rapidly growing population and increasing industry, along with weak regulatory measures, have led to uncontrolled waste and industrial discharge in coastal areas (<xref ref-type="bibr" rid="B24">Baqar et&#xa0;al., 2017</xref>). Industries, such as the Port Qasim, Korangi Power Plant, and Hub Power Plant (HUBCO) in Gaddani, often use seawater to cool heavy machinery and discharge warm salty water into adjacent coastal areas (<xref ref-type="bibr" rid="B144">Ullah et&#xa0;al., 2016</xref>). These discharges add a range of pollutants and excess nutrients to marine ecosystems, resulting in decreased water quality and increased turbidity, which has severe consequences for coral reefs, making it difficult for them to survive (<xref ref-type="bibr" rid="B3">Ahmad, 1997</xref>).</p>
<p>In addition to eutrophication, plastic pollution is a growing and concerning issue in the coastal areas of Pakistan and has a greater impact on the reefs and other marine ecosystems (<xref ref-type="bibr" rid="B119">Qaimkhani, 2018</xref>; <xref ref-type="bibr" rid="B127">Saher et&#xa0;al., 2019</xref>). Among South Asian countries, Pakistan has the highest percentage of mismanaged plastic (<xref ref-type="bibr" rid="B5">Ahmad et&#xa0;al., 2023</xref>). But there are limited studies regarding plastic pollution in coastal areas of Pakistan (<xref ref-type="bibr" rid="B120">Qari and Shaffat, 2015</xref>; <xref ref-type="bibr" rid="B42">Chaudhry, 2018</xref>; <xref ref-type="bibr" rid="B106">Neelam et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B119">Qaimkhani, 2018</xref>; <xref ref-type="bibr" rid="B5">Ahmad et&#xa0;al., 2023</xref>). However, a study conducted by (<xref ref-type="bibr" rid="B120">Qari and Shaffat, 2015</xref>) investigated the distribution and abundance of marine debris along the coast of Karachi and revealed that plastic is more dominant than other debris. In a recent study, <xref ref-type="bibr" rid="B5">Ahmad et&#xa0;al. (2023)</xref> asserted that microplastic pollution is emerging as a significant environmental concern in the coastal regions of Pakistan, exerting considerable impact on marine ecosystems.</p>
</sec>
<sec id="s4_3">
<label>4.3</label>
<title>Coastal tourism</title>
<p>Tourism in Pakistan&#x2019;s coastal areas has increased in recent years, with a diverse range of tourism attractions for climatic conditions, high mountain ranges, immense deserts, vast delta plains, and an amazing coastline (<xref ref-type="bibr" rid="B20">Arshad et&#xa0;al., 2018</xref>; <xref ref-type="bibr" rid="B131">Shahzad, 2020</xref>; <xref ref-type="bibr" rid="B6">Ahmed et&#xa0;al., 2022</xref>). These coastal regions are reflected globally, as maritime tourism is of important economic importance for various countries (<xref ref-type="bibr" rid="B48">Diakomihalis, 2007</xref>; <xref ref-type="bibr" rid="B71">Karani and Failler, 2020</xref>; <xref ref-type="bibr" rid="B87">Mart&#xed;nez V&#xe1;zquez et&#xa0;al., 2021</xref>). <xref ref-type="bibr" rid="B142">Ullah et&#xa0;al. (2010)</xref> highlighted that the tourism is currently playing a key role in strengthening Pakistan&#x2019;s shrinking economy. According to <xref ref-type="bibr" rid="B108">Nigar (2018)</xref> in 2015, the travel and tourism sector directly contributed $328.3 million to Pakistan&#x2019;s GDP, representing 2.8% of the total GDP. Realizing the importance of tourism in Pakistan&#x2019;s coastal zone, the government upgraded it to the status of an industry in 1989, highlighting its importance in promoting it to grow but has never been taken seriously as part of the overall tourism planning and development process (<xref ref-type="bibr" rid="B143">Ullah et&#xa0;al., 2018</xref>). There are approximately 28 beaches along its coastline, with most of them located in Sindh (20 beaches) and the remaining (8 beaches) in Balochistan (<xref ref-type="bibr" rid="B131">Shahzad, 2020</xref>), where coral reefs are scattered around the nearby waters. However, unsustainable tourism activities in Pakistan, such as boat anchoring and visitor snorkeling, pose direct damage by breaking or stepping on corals as well as indirect hazards to the reef ecosystem due to water pollution from shipping oil and visitor sunscreen (<xref ref-type="bibr" rid="B135">Siddiqui et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B117">Pakistan, 2016</xref>). Additionally, pollution from tourist-related actions such as litter, sewage, and boat-related oil spills can contaminate coastal waters and negatively impact marine ecosystems (<xref ref-type="bibr" rid="B119">Qaimkhani, 2018</xref>; <xref ref-type="bibr" rid="B55">Faran and Ejaz, 2022</xref>). Uncontrolled recreational activities such as snorkeling, scuba diving, and water sports can also harm coastal ecosystems with tourists potentially causing unintentional harm to coral reefs and disrupting delicate habitats, resulting in coral bleaching and decreased fish populations, with long-term consequences for the region&#x2019;s biodiversity (<xref ref-type="bibr" rid="B130">Sayied, 2007</xref>; <xref ref-type="bibr" rid="B119">Qaimkhani, 2018</xref>; <xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>). According to <xref ref-type="bibr" rid="B136">Siriwong et&#xa0;al. (2018)</xref> tourism has had terrible impacts on the marine ecosystems and coral reefs that are valuable to the tourism industry, have suffered severe consequences because of misuse by human activities such as sedimentation, habitat loss, waste dumping, and decreased light, endangering their health and existence.</p>
</sec>
<sec id="s4_4">
<label>4.4</label>
<title>Climate change and other challenges</title>
<p>Over recent years, South Asia has faced several devastating events due to the rapid increase of climate change (<xref ref-type="bibr" rid="B124">Rajasuriya et&#xa0;al., 2002</xref>). These effects include an increase in extreme weather events, rising temperatures, and sea-level rise, all of which have had severe impacts on marine ecosystems in the region. In Pakistan, the situation is particularly alarming (<xref ref-type="bibr" rid="B79">Khan et&#xa0;al., 2016</xref>). The increasing sea temperatures and ocean acidification, along with changing ocean currents, are profoundly affecting marine life (<xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). The increase in sea temperatures is leading to coral bleaching, which causing coral reefs to lose their vitality, disrupting the intricate symbiotic relationships they support and leading to the degradation of these vital ecosystems (<xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). The rise in sea levels and alterations in ocean currents further complicate the issue, influencing marine species distribution and abundance and potentially causing significant ecological shifts (<xref ref-type="bibr" rid="B83">Mahmood and Hassan, 2022</xref>). The National Institute of Oceanography (NIO) in Karachi reports a sea-level rise of approximately 1.1 millimeters per year in Pakistan&#x2019;s coastal region, which aligns with global trends and highlights the significant regional impacts of climate change observed over the past two decades (<ext-link ext-link-type="uri" xlink:href="https://niopk.gov.pk/">https://niopk.gov.pk/</ext-link>). For further studies related to climate change readers are directed to (<xref ref-type="bibr" rid="B56">Farooqi et&#xa0;al., 2005</xref>; <xref ref-type="bibr" rid="B101">Mustafa, 2011</xref>; <xref ref-type="bibr" rid="B83">Mahmood and Hassan, 2022</xref>; <xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). Despite these pressing issues, climate change-related studies investigating these effects on Pakistan&#x2019;s coastal water are still very limited (<xref ref-type="bibr" rid="B121">Qureshi, 2005</xref>). This gap in research is critical given that Pakistan is ranked fifth on the Global Climate Risk Index for vulnerability to climate change impacts (<xref ref-type="bibr" rid="B128">Salik et&#xa0;al., 2015</xref>).</p>
<p>In addition, the sedimentation or siltation of sand and silt along Pakistan&#x2019;s coast, primarily caused by the Indus River&#x2019;s drainage system, has a significant impact on the marine ecosystems (<xref ref-type="bibr" rid="B27">Beg, 1995</xref>; <xref ref-type="bibr" rid="B76">Khan et&#xa0;al., 2021</xref>). Apart from free-living species, once-settled corals are sessile organisms (<xref ref-type="bibr" rid="B38">Chadwick, 1988</xref>; <xref ref-type="bibr" rid="B39">Chadwick-Furman and Loya, 1992</xref>; <xref ref-type="bibr" rid="B118">Pedersen et&#xa0;al., 2019</xref>), rely on their development form and physiological adaptations to regulate their interactions with the environment because they can&#x2019;t move away from unfavorable conditions (<xref ref-type="bibr" rid="B98">Muller-Parker et&#xa0;al., 2015</xref>). Reef-building corals rely on symbiotic, unicellular algae known as zooxanthellae, which live as symbiotic inside the coral tissue, primarily in the gastrodermis, and through photosynthesis produce most of the coral&#x2019;s energy requirements, because of this symbiotic interaction, most corals rely on light for survival (<xref ref-type="bibr" rid="B61">Harrison and Booth, 2007</xref>; <xref ref-type="bibr" rid="B98">Muller-Parker et&#xa0;al., 2015</xref>; <xref ref-type="bibr" rid="B52">El-Naggar, 2020</xref>). Turbidity and sedimentation caused by coastal building and dredging are the major causes of diminished light penetration and sediment deposition on the coral&#x2019;s surface, making it difficult for the coral&#x2019;s symbiotic zooxanthellae to capture enough light for photosynthesis. They also complicate the coral&#x2019;s feeding when it is buried in sediment, as well as dealing with suspended sediments, particularly fine-grained, imposes additional energy costs on the coral due to the need for mucus production, sediment clearance, and impaired feeding, all of which harm zooxanthellae (<xref ref-type="bibr" rid="B53">Erftemeijer et&#xa0;al., 2012</xref>). Turbidity decreases ambient photosynthetically active radiation (PAR), which reduces zooxanthellae productivity and can lead to starvation. Sediment settling on coral tissue produces additional shadowing and smothering, contributing to a further decline in zooxanthellae photosynthetic activity and even coral bleaching (<xref ref-type="bibr" rid="B31">Borell and Bischof, 2008</xref>; <xref ref-type="bibr" rid="B57">Freitas et&#xa0;al., 2019</xref>).</p>
</sec>
</sec>
<sec id="s5">
<label>5</label>
<title>Conservation status and management recommendations</title>
<p>Pakistan is a federal state with three tiers of government-local, provincial, and central. They are working by established laws and regulations but the federal government taking the responsibility for the overall strategy (<xref ref-type="bibr" rid="B147">Ullah et&#xa0;al., 2022</xref>). In this context various government departments, community-based initiatives, and NGOs in Pakistan collaborate directly with coastal communities to protect coastal habitats (<xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref>). These attempts usually entail joint efforts with government departments, research institutions, and organizations with these three tiers of government to achieve their goals of conserving coastal ecosystems. There is potential legislation to protect marine life (Biodiversity Action Plan 1997, Pakistan Environmental Protection Act 1995, Environmental Protection Ordinance 1983, and the Wildlife Protection Ordinance 1972) (<xref ref-type="bibr" rid="B124">Rajasuriya et&#xa0;al., 2002</xref>). Despite this, clear policies for managing coastal and marine resources have not yet been fully covered, and the existing policies, laws, and legislation in the country remain confusing and complex (<xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). These factors have led to uncertainty among organizations about involving coastal communities in the planning and development process. The current laws and regulations governing coastal and marine areas in Pakistan lack an integrated management framework and only provide guidelines for environmentally friendly economic activities. However, the present situation of coral reefs and other ecosystems shows that the conservation of coral reefs requires a paradigm shift in planning and policy frameworks, management practices, research with tangible outcomes, the active involvement and coordination of stakeholders, capacity building, and a strong commitment from all relevant government bodies (<xref ref-type="fig" rid="f5">
<bold>Figure&#xa0;5</bold>
</xref>).</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Comparison of coral reef conservation policies and practices in Pakistan with natural and anthropogenic threats.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g005.tif"/>
</fig>
<sec id="s5_1">
<label>5.1</label>
<title>Strengthen legislation and enforcement</title>
<p>Pakistan has implemented conservation policies and enacted laws, strategies, and policies for the conservation of coral reefs and other marine ecosystems. However, there is still a need for substantial efforts in this direction. The Karachi harbor area is a stark example of one of the most polluted areas for oil along the coast of Pakistan (<xref ref-type="bibr" rid="B133">Siddique et&#xa0;al., 2009</xref>). The harbor covers approximately 25 square kilometers, receives approximately 50 billion cubic meters of seawater per tidal cycle, and is visited by 3,000 to 4,000 ships annually (<xref ref-type="bibr" rid="B130">Sayied, 2007</xref>). During regular operations, ships discharge bilge water, oily ballast water, and cargo tank washings, with large amounts of pollution stemming from oily spills from both oil tankers and general cargo vessels (<xref ref-type="bibr" rid="B130">Sayied, 2007</xref>; <xref ref-type="bibr" rid="B134">Siddiqui and Amir, 2010</xref>). Although laws and regulations designed to control pollution are present, they are rarely enforced or implemented (<xref ref-type="bibr" rid="B130">Sayied, 2007</xref>). As highlighted by <xref ref-type="bibr" rid="B99">Mumtaz et&#xa0;al. (2023)</xref> poor governance has led to the degradation of marine ecosystems in coastal areas of Pakistan. Poor governance refers to the system and processes by which several actors (e.g., government, nongovernmental organizations, and the private sector) share authority and make decisions. This encompasses formal laws and regulations as well as more dispersed components such as media, trade, religion, and public opinion (<xref ref-type="bibr" rid="B99">Mumtaz et&#xa0;al., 2023</xref>). The absence of specific maritime laws addressing governance in Pakistan complicates the implementation of coastal and marine planning (<xref ref-type="bibr" rid="B146">Ullah et&#xa0;al., 2021</xref>). Despite existing laws, no legislation comprehensively covers the issues of the country&#x2019;s coastal areas (<xref ref-type="bibr" rid="B146">Ullah et&#xa0;al., 2021</xref>). To enhance Pakistan&#x2019;s coastal and marine conservation efforts, the country must adopt and adhere to effective guidelines, laws, and policy frameworks similar to those implemented by China. China&#x2019;s governance model, which emphasizes rule by law, has fostered new initiatives and innovations in the sustainable development of coastal and marine resources (<xref ref-type="bibr" rid="B43">Chen and Pearson, 2015</xref>). Several laws and regulations have been enacted to oversee administrative jurisdiction rights pertaining to coastal and marine areas. The two most significant pieces of legislation, the Marine Environment Protection Law 1999 and the LAUSA 2001 have been enacted to regulate the marine environment and to utilize marine resources rationally and sustainably (<xref ref-type="bibr" rid="B74">Keyuan, 2001</xref>; <xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). These two laws are considered as important and supportive legislation for China&#x2019;s marine functional zoning (MFZ), covering all areas of environmental conservation and sustainable use of coastal and marine natural resources (<xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). The effective enforcement of these two laws has bolstered and enhanced China&#x2019;s sea use authorization system, marine environment protection measures, marine functional zoning framework, and marine legal infrastructure (<xref ref-type="bibr" rid="B81">Li, 2006</xref>; <xref ref-type="bibr" rid="B37">Cao and Wong, 2007</xref>; <xref ref-type="bibr" rid="B82">Lu et&#xa0;al., 2015</xref>). The implementation of laws within the MFZ schemes delineates the administrative responsibilities of the authorities engaged in managing coastal and marine zones. For Pakistan to achieve similar success, it must create specific policy frameworks/laws at various administrative levels and national, provincial, district, and tehsil levels. These frameworks require well-developed development strategies, good governance, improved stakeholder coordination, and continuous restoration and rehabilitation initiatives (<xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>). Poor governance, inadequate planning, implementation of development plans, and a lack of coordination among stakeholders within the current framework may result in irreversible loss and degradation of coral reefs and other ecosystems in Pakistan (<xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>).</p>
<p>Moreover, effective coral reef conservation laws require adequate financial resources to support these invaluable marine ecosystems (<xref ref-type="bibr" rid="B49">Dixit et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). An effective approach to raising funds involves imposing an environmental surcharge on commercial entities, shipbreaking operations, cargo vessels, ferry services, industrial units, hotels, and others that utilize coral reef areas for their operations or contribute to marine pollution (<xref ref-type="bibr" rid="B36">Byrnes and Dunn, 2020</xref>). Therefore, the government should provide incentives and tax breaks to industries that adopt pollution prevention measures. Funding should be allocated for safeguarding and protecting critical habitats and providing training and capacity building for the staff engaged in the conservation and management of coral reefs. The country lacks a regulatory framework that would support an integrated management strategy to properly manage coastal and marine resources (<xref ref-type="bibr" rid="B146">Ullah et&#xa0;al., 2021</xref>). In Pakistan, however, plans and policies are developed at the national level and implemented by lower-level governments (<xref ref-type="bibr" rid="B116">Pakistan, 2014</xref>). These plans also fulfil sector-specific objectives and emphasize resource marketization and privatization, resulting in a prevalent paradigm of resource empowerment or ocean governance (<xref ref-type="bibr" rid="B116">Pakistan, 2014</xref>). Therefore, Pakistan should meet the needs of marine environment conservation and resource usage. Implementing such a legal framework is not only critical to the sustainable management of marine biodiversity, but also contributes to ensuring that conservation efforts are holistic, consistent, and strong in the long term. Pakistan may adopt global best practices to emulate a functional conservation framework, especially from countries with successful records in marine conservation. For example, Australia&#x2019;s Great Barrier Reef Marine Park Authority (GBRMPA) suggested a plan for managing large-scale marine parks through legally defined governance approaches, stakeholder engagement, and adaptive management processes (<xref ref-type="bibr" rid="B54">Evans et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B138">Stephenson et&#xa0;al., 2023</xref>). Similar strategies, when implemented, could act as a roadmap to enable Pakistan to develop robust marine protected areas. Pakistan can enhance its conservation practices, protect its valuable coral reefs, and ensure the sustainable utilization of marine resources for generations by understanding global lessons and localizing strategies. Adopting this integrated approach is necessary not only to address prevailing issues but also to develop resilience against other potential threats to the ecosystem. <xref ref-type="fig" rid="f6">
<bold>Figure&#xa0;6</bold>
</xref> visually supports this discussion by illustrating the consequences of inadequate conservation laws and governance on reef systems. Showing the hierarchy of causes, global practices, proposed solutions, and implementation strategies. The arrows indicate the flow of influence and interactions between these elements, while the colors differentiate the categories for clarity.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Framework for restoring coral reefs: A guide to coral reef restoration (The referenced literature and <xref ref-type="table" rid="T1">
<bold>Table&#xa0;1</bold>
</xref> were used to create this figure).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-11-1466834-g006.tif"/>
</fig>
</sec>
<sec id="s5_2">
<label>5.2</label>
<title>Enhance protection awareness</title>
<p>Insufficient awareness of ecological and environmental issues globally contributes to the degradation of coastal ecosystems (<xref ref-type="bibr" rid="B73">Kautsky et&#xa0;al., 2000</xref>; <xref ref-type="bibr" rid="B58">Garcia and De Leiva Moreno, 2003</xref>; <xref ref-type="bibr" rid="B26">Bayraktarov et&#xa0;al., 2016</xref>). The deterioration of marine ecosystems worldwide requires urgent global management strategies to safeguard their resilience (<xref ref-type="bibr" rid="B90">Mccook et&#xa0;al., 2009</xref>).</p>
<p>The coastal wetlands of Pakistan include a wide range of ecosystems such as coral reefs, mangroves, estuaries, and beaches (<xref ref-type="bibr" rid="B41">Chaudhry, 2010</xref>). Despite the ecological and economic importance of marine biodiversity protection, awareness in Pakistan remains critically inadequate due to weak protection awareness (<xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). However, in Pakistan limited legislative improvements, rapid economic growth, and urbanization continue to overwhelm regulatory institutions, hampered by a lack of public awareness or concern regarding environmental degradation (<xref ref-type="bibr" rid="B129">Sardar, 2012</xref>; <xref ref-type="bibr" rid="B102">Mustafa et&#xa0;al., 2013</xref>). As a result, one of the serious threats to the coral reef ecosystem in Pakistan is a lack of awareness and understanding among the general population (<xref ref-type="bibr" rid="B124">Rajasuriya et&#xa0;al., 2002</xref>; <xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). People who live near these areas are unaware of the importance of coral reefs in supporting coastal ecosystems (<xref ref-type="bibr" rid="B41">Chaudhry, 2010</xref>). Consequently, there is limited public support for conservation efforts (<xref ref-type="bibr" rid="B144">Ullah et&#xa0;al., 2016</xref>). The large commercial tours contribute to environmental litter, with items such as food and plastic being thrown into the water (<xref ref-type="bibr" rid="B144">Ullah et&#xa0;al., 2016</xref>). large resources allocated to monitoring coral reefs often render these ecosystems vulnerable to illegal activities like destructive fishing practices, overfishing, and coral mining, particularly along the Karachi coast where an increasing number of unauthorized fishing villages further exacerbate the situation (<xref ref-type="bibr" rid="B122">Qureshi, 2011</xref>; <xref ref-type="bibr" rid="B91">Mehak et&#xa0;al., 2023</xref>). This discussion emphasizes the need for the conservation of several species and habitats. However, between 2013 and 2016, there was a ban on fishing in the area from May to August, which was initially imposed by the Karachi Fisheries Harbor Authority to protect the population during the breeding season. However, the ban proved ineffective due to weak awareness among local communities and little logistical control over the area (<xref ref-type="bibr" rid="B93">Moazzam and Osmany, 2021</xref>; <xref ref-type="bibr" rid="B113">Osmany and Moazzam, 2022</xref>). This issue requires delicate handling, necessitating the exploration of alternative employment opportunities or income sources for residents. This can be achieved through the development of small-scale cottage industries or facilitating access to financing from banks for small businesses. Implementing a blanket ban may not always be feasible, especially in areas heavily dependent on fisheries. Central to this strategy will be the empowerment of communities whose lives are inextricably linked to the resources in question. <xref ref-type="bibr" rid="B2">Aguero and Costello (1986)</xref> observe that problems related to fisheries are rarely resolved by focusing solely on fishery management proposes. <xref ref-type="bibr" rid="B114">Ostrom (1990)</xref> has demonstrated that the conservation of natural resources is facilitated when local communities actively participate in the process. Additionally, <xref ref-type="bibr" rid="B34">Bromley (1992)</xref> argues that cooperation among people in conserving an open-access resource (OAR) is facilitated when they possess the capability to devise, oversee, and enforce regulations governing the utilization and allocation of natural resources by members.</p>
<p>To address these threats and enhance the protection and awareness of coral reefs in Pakistan&#x2019;s coastal zone, a few sites are recommended for the designation of MPAs. The most important site may be Astola Island. However, Pakistan has designated Astola Island as an MPA (<xref ref-type="bibr" rid="B135">Siddiqui et&#xa0;al., 2008</xref>). Additionally, the IUCN has recommended Churna Island as one of the areas that should receive this protection (<xref ref-type="bibr" rid="B63">Hoon et&#xa0;al., 1997</xref>; <xref ref-type="bibr" rid="B119">Qaimkhani, 2018</xref>). While there might be some challenges for fishermen because of imposing protected areas. However, there might be some challenges for fishermen because of the need to establish protected areas. However, there are several advantages to the establishment of MPAs at various levels, such as marine ecosystem conservation, contributions to sustainable tourism, genetic diversity maintenance, potential enhancement of ecosystem-based fisheries management (especially in coastal areas crucial for breeding and spawning of economically valuable species), and preservation of prime locations for recreational activities such as SCUBA diving. Designating the MPA status would impose restrictions within a specified radius of the island, providing a safe breeding habitat for fish. Commercial activities should also be limited to reduce pollution and damage. Officially designating protected status could also raise public awareness of the island&#x2019;s importance for biodiversity, making it more challenging for companies to disregard necessary protection. Additionally, various natural and anthropogenic pressures on Pakistan&#x2019;s coral reefs can be reduced to the greatest extent possible by diverse management practices, development projects, legislative regulations, awareness-raising initiatives, and community and NGO involvement. Furthermore, strict environmental regulation enforcement is essential to counteract illegal activities that destroy coral reefs. Concerned departments should conduct comprehensive investigations of all coastal areas and collect data to determine the sites with the most potential for coral species rehabilitation, particularly for the restoration of extinct species. They should engage the assistance of local governments, local communities, volunteers, NGOs, religious leaders, elders, and the media to ensure that each sector of society contributes to the success of rehabilitation efforts and the efficient management and conservation of coral reefs.</p>
</sec>
<sec id="s5_3">
<label>5.3</label>
<title>Provide scientific support</title>
<p>In recent years, various countries have made significant progress globally in enhancing their scientific knowledge of coastal ecosystems. Consequently, the conservation of coastal ecosystems in Pakistan needs strong scientific support (<xref ref-type="bibr" rid="B135">Siddiqui et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B145">Ullah et&#xa0;al., 2017</xref>). In Pakistan, the continuous decline of the coral reef ecosystem further highlights the need for comprehensive scientific investigations (<xref ref-type="bibr" rid="B14">Ali et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B10">Ali and Hussain, 2023</xref>). Research on coral reefs has acquired critical significance globally due to their high biodiversity and aesthetic beauty (<xref ref-type="bibr" rid="B33">Brodie and Waterhouse, 2012</xref>; <xref ref-type="bibr" rid="B100">Muruganandam et&#xa0;al., 2023</xref>). However, it is necessary to conduct a comprehensive scientific analysis of the threats confronting coral reefs in Pakistan. Assessing the extent of these unraveling and threats to their interconnections enables researchers to address the most pressing issues. For example, <xref ref-type="bibr" rid="B11">Ali et&#xa0;al. (2014)</xref> highlight the impacts of overfishing, destructive fishing practices, and pollution in the country&#x2019;s coral reefs. Recommended management in response to these threats includes completing baseline assessments and coral reef research, establishing MPAs, and developing management initiatives and laws. <xref ref-type="bibr" rid="B12">Ali et&#xa0;al. (2017)</xref> investigated the factors influencing coral growth, focusing on the challenges posed by sedimentation and rising sea temperatures. The authors suggested the implementation of laws to manage coastal development and water quality. <xref ref-type="bibr" rid="B66">Iqbal et&#xa0;al. (2018)</xref> studied is about the coral reef responses to climate change. Their recommended proposed management approach involves remote sensing to monitor the impacts of climate change on the coral reef ecosystems. <xref ref-type="bibr" rid="B13">Ali et&#xa0;al. (2020)</xref> investigated the biodiversity reduction and faunal turnover in coral reefs. Their study suggested identifying and mitigating specific coral reef stresses. In addition, <xref ref-type="bibr" rid="B14">Ali et&#xa0;al. (2021)</xref> focused on unsustainable tourism practices and inadequate community involvement and awareness. Their study suggested reef restoration efforts and conservation initiatives.</p>
<p>These studies significantly contribute to a comprehensive understanding of the challenges confronting coral reefs and encourage conservation strategies. The scientific research, monitoring actions, and engagement community have emerged as a consistent thread, reflecting a concerted effort to increase the resilience and long-term survival of coral reefs. For example, China, a country with an extensive coastline (South China Sea), faces similar challenges regarding the conservation of its coastal environment (<xref ref-type="bibr" rid="B64">Huang et&#xa0;al., 2016</xref>). The South China Sea, located in the central Indo-Pacific region, is a large semi enclosed marine environment that supports diverse coral reef ecosystems (<xref ref-type="bibr" rid="B95">Morton and Blackmore, 2001</xref>). In the South China Sea, scientific research strongly supports the conservation of coral reefs, underlying their critical role in sustaining marine biodiversity, fisheries, and other ecosystem services. Various studies have highlighted the ecological importance and long-term conservation of this fragile ecosystem to overcome the impacts of climate change, overfishing, pollution, and other threats (<xref ref-type="bibr" rid="B44">Chou et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B18">Arai, 2015</xref>; <xref ref-type="bibr" rid="B64">Huang et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B151">Zhao et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B22">Asner et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B152">Zhao et&#xa0;al., 2019</xref>). Similarly, scientific research should be considered in Pakistan to enhance the effective conservation of coral reefs, following the example set by China. To overcome the challenges confronting these ecosystems, collaboration among managers and experts from various fields, such as marine geology, marine biology, oceanography, ecology, and local knowledge, is needed. Scientific efforts should be made to understand the distinctive dynamics of coral reefs in coastal areas of Pakistan. By collaborating with managers and experts from different backgrounds, Pakistan can gain a holistic perspective on the threats affecting its coral reef ecosystem.</p>
<p>Furthermore, collaborative research involving local researchers such as (<xref ref-type="bibr" rid="B11">Ali et&#xa0;al., 2014</xref>) and foreign researchers is needed to collect accurate and current data on coral distribution, variety, abundance, and threats in the coastal area of Pakistan.</p>
</sec>
</sec>
<sec id="s6" sec-type="conclusions">
<label>6</label>
<title>Conclusion</title>
<p>The threats and conservation efforts to coral reefs in Pakistan have been thoroughly examined in this review. Coral reefs are an important coastal resource in Pakistan, contributing significantly to the country&#x2019;s rapid social and economic growth. However, these fragile ecosystems have been disturbed by a variety of human activities with high frequency and intensity. The main anthropogenic threats to a country&#x2019;s coral ecosystems are overfishing, pollution, and tourism. The effective protection and management of coral reef ecosystems are emergent for both local and central governments. The difficulties lie in the lack of protection awareness, targeted law, and adequate scientific basis. The promotion of national awareness of environmental protection is necessary and critical for reducing destructive activities and increasing conservation activities. Various targeted and effective protection measures could be implemented according to environmental policy and regulations; for example, the discharge of untreated sewage and destructive fishing are forbidden. In addition, rich and accurate information on coral reef ecosystems could provide a solid scientific foundation for the rapid understanding and management of coral reefs. To effectively address the conservation challenges affecting the country&#x2019;s coral reefs, government support, researchers, and academia are needed. Furthermore, it is urgent to establish MPAs along Pakistan&#x2019;s coastline to protect these ecosystems from further degradation.</p>
</sec>
</body>
<back>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>IA: Methodology, Writing &#x2013; original draft, Software. PG: Writing &#x2013; review &amp; editing, Conceptualization. M-XZ: Writing &#x2013; review &amp; editing, Supervision, Investigation, Funding acquisition. YZ: Writing &#x2013; review &amp; editing, Software. X-YZ: Writing &#x2013; review &amp; editing, Visualization. S-QZ: Writing &#x2013; review &amp; editing, Methodology. J-WQ: Writing &#x2013; review &amp; editing, Formal Analysis. QS: Writing &#x2013; review &amp; editing, Conceptualization. H-QY:&#xa0;Writing &#x2013; review &amp; editing, Validation. S-CT: Writing &#x2013; review &amp; editing, Project administration. L-JX: Writing &#x2013; review &amp; editing, Data curation.</p>
</sec>
<sec id="s8" 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. This research was financially supported by the National Key R&amp;D Program of China (Grant No. 2021YFC3100600), National Natural Science Foundation of China project (Grant Nos. 41876132,41806139 and 42376165), Hainan Province Science and Technology Special Fund (Grant No. ZDYF2022SHFZ072), and the Open Research Fund Program of Guangxi Key Lab of Mangrove Conservation and Utilization (Grant No. GKLMC-201904).</p>
</sec>
<sec id="s9" 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="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>
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