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<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Mar. Sci.</journal-id>
<journal-title-group>
<journal-title>Frontiers in Marine Science</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Mar. Sci.</abbrev-journal-title>
</journal-title-group>
<issn pub-type="epub">2296-7745</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
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<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2025.1663369</article-id>
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<article-categories>
<subj-group subj-group-type="heading">
<subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>CoDeMap: a classification scheme for benthic habitats from the coast to the deep sea in the Mediterranean and Black Sea</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author">
<name><surname>Grande</surname><given-names>Valentina</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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<name><surname>Angeletti</surname><given-names>Lorenzo</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<xref ref-type="author-notes" rid="fn003"><sup>&#x2020;</sup></xref>
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<contrib contrib-type="author" corresp="yes">
<name><surname>Prampolini</surname><given-names>Mariacristina</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="corresp" rid="c001"><sup>*</sup></xref>
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<contrib contrib-type="author">
<name><surname>Castellan</surname><given-names>Giorgio</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
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<contrib contrib-type="author">
<name><surname>Dalla Valle</surname><given-names>Giacomo</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3154134/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; original draft" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-original-draft/">Writing &#x2013; original draft</role>
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<name><surname>Fraschetti</surname><given-names>Simonetta</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff4"><sup>4</sup></xref>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
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<contrib contrib-type="author">
<name><surname>Basso</surname><given-names>Daniela</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
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<contrib contrib-type="author">
<name><surname>Berov</surname><given-names>Dimitar</given-names></name>
<xref ref-type="aff" rid="aff7"><sup>7</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3228640/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="validation" vocab-term-identifier="https://credit.niso.org/contributor-roles/validation/">Validation</role>
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<contrib contrib-type="author">
<name><surname>Bracchi</surname><given-names>Valentina A.</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/1377151/overview"/>
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<contrib contrib-type="author">
<name><surname>Cardone</surname><given-names>Frine</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<xref ref-type="aff" rid="aff8"><sup>8</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/529170/overview"/>
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<contrib contrib-type="author">
<name><surname>Chimienti</surname><given-names>Giovanni</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff9"><sup>9</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/846592/overview"/>
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<contrib contrib-type="author">
<name><surname>Falace</surname><given-names>Annalisa</given-names></name>
<xref ref-type="aff" rid="aff10"><sup>10</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/804753/overview"/>
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<contrib contrib-type="author">
<name><surname>Galil</surname><given-names>Bella</given-names></name>
<xref ref-type="aff" rid="aff11"><sup>11</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/3228710/overview"/>
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<contrib contrib-type="author">
<name><surname>Mastrototaro</surname><given-names>Francesco</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff9"><sup>9</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/2038286/overview"/>
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<contrib contrib-type="author">
<name><surname>Salomidi</surname><given-names>Maria</given-names></name>
<xref ref-type="aff" rid="aff12"><sup>12</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/156806/overview"/>
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<contrib contrib-type="author">
<name><surname>Savini</surname><given-names>Alessandra</given-names></name>
<xref ref-type="aff" rid="aff5"><sup>5</sup></xref>
<xref ref-type="aff" rid="aff6"><sup>6</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/476389/overview"/>
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<contrib contrib-type="author">
<name><surname>Schembri</surname><given-names>Patrick J.</given-names></name>
<xref ref-type="aff" rid="aff13"><sup>13</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/1387964/overview"/>
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<contrib contrib-type="author">
<name><surname>Todorova</surname><given-names>Valentina</given-names></name>
<xref ref-type="aff" rid="aff14"><sup>14</sup></xref>
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<contrib contrib-type="author">
<name><surname>Taviani</surname><given-names>Marco</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="aff" rid="aff8"><sup>8</sup></xref>
<xref ref-type="author-notes" rid="fn004"><sup>&#x2021;</sup></xref>
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<name><surname>Foglini</surname><given-names>Federica</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="fn004"><sup>&#x2021;</sup></xref>
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<aff id="aff1"><label>1</label><institution>National Research Council-Institute of Marine Sciences (CNR-ISMAR)</institution>, <city>Bologna</city>, <country country="it">Italy</country></aff>
<aff id="aff2"><label>2</label><institution>National Biodiversity Future Center (NBFC)</institution>, <city>Palermo</city>, <country country="it">Italy</country></aff>
<aff id="aff3"><label>3</label><institution>National Research Council-Institute for Marine Biological Resources and Biotechnology (CNR-IRBIM)</institution>, <city>Ancona</city>, <country country="it">Italy</country></aff>
<aff id="aff4"><label>4</label><institution>Department of Biology, University of Naples Federico II</institution>, <city>Naples</city>, <country country="it">Italy</country></aff>
<aff id="aff5"><label>5</label><institution>Consorzio Nazionale Interuniversitario per le Scienze del Mare (CoNISMa)</institution>, <city>Roma</city>, <country country="it">Italy</country></aff>
<aff id="aff6"><label>6</label><institution>Department of Earth and Environmental Sciences, University of Milano-Bicocca</institution>, <city>Milano</city>, <country country="it">Italy</country></aff>
<aff id="aff7"><label>7</label><institution>Institute of Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences</institution>, <city>Sofia</city>, <country country="bg">Bulgaria</country></aff>
<aff id="aff8"><label>8</label><institution>Stazione Zoologica &#x201c;Anton Dohrn&#x201d;</institution>, <city>Napoli</city>, <country country="it">Italy</country></aff>
<aff id="aff9"><label>9</label><institution>Department of Biosciences, Biotechnologies and Environment, University of Bari Aldo Moro</institution>, <city>Bari</city>, <country country="it">Italy</country></aff>
<aff id="aff10"><label>10</label><institution>Department of Life Sciences, University of Trieste</institution>, <city>Trieste</city>, <country country="it">Italy</country></aff>
<aff id="aff11"><label>11</label><institution>The Steinhardt Museum of Natural History, Tel Aviv University</institution>, <city>Tel Aviv</city>, <country country="il">Israel</country></aff>
<aff id="aff12"><label>12</label><institution>Institute of Oceanography, Hellenic Centre for Marine Research</institution>, <city>Anavyssos</city>, <country country="gr">Greece</country></aff>
<aff id="aff13"><label>13</label><institution>Faculty of Science, University of Malta</institution>, <city>Msida</city>, <country country="mt">Malta</country></aff>
<aff id="aff14"><label>14</label><institution>Bulgarian Academy of Sciences, Institute of Oceanology</institution>, <city>Varna</city>, <country country="bg">Bulgaria</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Mariacristina Prampolini, <email xlink:href="mailto:mariacristina.prampolini@cnr.it">mariacristina.prampolini@cnr.it</email></corresp>
<fn fn-type="other" id="fn003">
<label>&#x2020;</label>
<p>These authors share first authorship</p></fn>
<fn fn-type="other" id="fn004">
<label>&#x2021;</label>
<p>These authors share last authorship</p></fn>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2025-11-06">
<day>06</day>
<month>11</month>
<year>2025</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2025</year>
</pub-date>
<volume>12</volume>
<elocation-id>1663369</elocation-id>
<history>
<date date-type="received">
<day>10</day>
<month>07</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>29</day>
<month>09</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Grande, Angeletti, Prampolini, Castellan, Dalla Valle, Fraschetti, Basso, Berov, Bracchi, Cardone, Chimienti, Falace, Galil, Mastrototaro, Salomidi, Savini, Schembri, Todorova, Taviani and Foglini.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Grande, Angeletti, Prampolini, Castellan, Dalla Valle, Fraschetti, Basso, Berov, Bracchi, Cardone, Chimienti, Falace, Galil, Mastrototaro, Salomidi, Savini, Schembri, Todorova, Taviani and Foglini</copyright-holder>
<license>
<ali:license_ref start_date="2025-11-06">https://creativecommons.org/licenses/by/4.0/</ali:license_ref>
<license-p>This is an open-access article distributed under the terms of the <ext-link ext-link-type="uri" xlink:href="https://creativecommons.org/licenses/by/4.0/">Creative Commons Attribution License (CC BY)</ext-link>. 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.</license-p>
</license>
</permissions>
<abstract>
<p>The spatial representation of benthic habitats is essential across various applications, such as biodiversity monitoring, ecosystem management and conservation, and maritime spatial planning. In this context, classification schemes provide a universally understandable framework to characterize and chart the seafloor. This work introduces the Coast to Deep Mapping (CoDeMap) classification scheme for benthic habitats from the coast to the deep-sea environments. It consists of three main components (Morphology, Substrate and Biology) and it is conceived as a practical tool for users from various backgrounds who need to organize and interpret marine observational data, as well as characterize and map seafloors. While primarily developed for the Mediterranean Sea and the Black Sea, CoDeMap serves as a foundational framework that can be adapted to address any current or future similar request worldwide.</p>
</abstract>
<kwd-group>
<kwd>benthic habitat mapping</kwd>
<kwd>morphology</kwd>
<kwd>substrate</kwd>
<kwd>biology</kwd>
<kwd>hierarchical classification scheme</kwd>
<kwd>GIS</kwd>
<kwd>Mediterranean Sea</kwd>
<kwd>Black Sea</kwd>
</kwd-group>
<funding-group>
<funding-statement>The author(s) declare financial support was received for the research and/or publication of this article. This paper is CNR-ISMAR-Bologna scientific contribution n. 2091 and is part of EU F.P. VII Project CoCoNet (contract no. 287844), the Italian Flagship Project RITMARE, LIFE DREAM Project (LIFE21-NAT-IT-LIFE-DREAM/101074547), REDRESS Project (project no. 101135492), and the National Biodiversity Future Center - NBFC (funded under the National Recovery and Resilience Plan (NRRP), Mission 4 Component 2 Investment 1.4 - Call for tender No. 3138 of 16 December 2021, rectified by Decree n.3175 of 18 December 2021 of Italian Ministry of University and Research funded by the European Union &#x2013; NextGenerationEU; Award Number: Project code CN_00000033, Concession Decree No. 1034 of 17 June 2022 adopted by the Italian Ministry of University and Research, CUP D33C22000960007).</funding-statement>
</funding-group>
<counts>
<fig-count count="8"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="66"/>
<page-count count="16"/>
<word-count count="6125"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Marine Ecosystem Ecology</meta-value>
</custom-meta>
</custom-meta-group>
</article-meta>
</front>
<body>
<sec id="s1" sec-type="intro">
<label>1</label>
<title>Introduction</title>
<p>The term &#x201c;habitat&#x201d; refers to the geographical, abiotic, and biotic characteristics of the environment where a species resides in any state of its life cycle. Habitat is an essential element of the seascape, frequently associated with diversity, functioning, and ecosystem services (<xref ref-type="bibr" rid="B57">Soko&#x142;owski et&#xa0;al., 2021</xref>). As a result, habitats became the primary classification unit in marine cartography and the focus of inventories, classification systems, and spatial mapping efforts (<xref ref-type="bibr" rid="B14">Coggan et&#xa0;al., 2007</xref>). In mapping, &#x201c;habitat&#x201d; is often used with a broader meaning and embraces more species, coming closer to the term &#x201c;biotope&#x201d;, i.e., the physical conditions in which a specific group of species lives (<xref ref-type="bibr" rid="B46">Montefalcone et&#xa0;al., 2021</xref>). <xref ref-type="bibr" rid="B45">Misiuk and Brown (2024)</xref> define benthic habitat mapping as &#x201c;a spatially continuous prediction of biological patterns on the seafloor,&#x201d; refining the earlier definition provided by <xref ref-type="bibr" rid="B10">Brown et&#xa0;al. (2011)</xref>, which described it as &#x201c;the use of spatially continuous environmental data sets to represent and predict biological patterns on the seafloor (whether continuous or discontinuous).&#x201d;</p>
<p>The spatial representation of the distribution and extent of physically distinct areas of the seafloor, which are linked to groups of species or communities that consistently coexist (<xref ref-type="bibr" rid="B35">Harris and Baker, 2020</xref>), is vital for several reasons. In fact, maps on the distribution of benthic habitats facilitate to:</p>
<list list-type="simple">
<list-item>
<p>- identify biodiversity hotspots and provide inventories of vulnerable species and ecosystems, and critical or sensitive areas (<xref ref-type="bibr" rid="B64">Vassallo et&#xa0;al., 2018</xref>);</p></list-item>
<list-item>
<p>- orient conservation actions by identifying priority areas for protection (<xref ref-type="bibr" rid="B2">Angeletti et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B65">Ware and Downie, 2020</xref>) and plan effective management strategies (<xref ref-type="bibr" rid="B24">Fraschetti et&#xa0;al., 2011</xref>);</p></list-item>
<list-item>
<p>- consider a habitat-based approach in policy support and decision-making processes (<xref ref-type="bibr" rid="B9">Bianchi et&#xa0;al., 2012</xref>; <xref ref-type="bibr" rid="B17">Danovaro et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B57">Soko&#x142;owski et&#xa0;al., 2021</xref>);</p></list-item>
<list-item>
<p>- meet the requirements of European directives and programs (<xref ref-type="bibr" rid="B56">Schiele et&#xa0;al., 2014</xref>), such as the Habitat Directive (92/43/EEC), the Marine Strategy Framework Directive (2008/56/EC), the Biodiversity Strategy for 2030 (EU, 2020), the Nature Restoration Law (EU Regulation 2022/869);</p></list-item>
<list-item>
<p>- monitor anthropogenic impacts, environmental status and trends (<xref ref-type="bibr" rid="B7">Bekkby et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B23">Enrichetti et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B28">Gerovasileiou et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B37">Holon et&#xa0;al., 2015</xref>);</p></list-item>
<list-item>
<p>- assess seafloor economic resources and quantify ecosystem services (<xref ref-type="bibr" rid="B13">Cogan et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B42">McQuaid et&#xa0;al., 2020</xref>);</p></list-item>
<list-item>
<p>- implement modeling approaches to predict areas suitable for species and communities and detect changes (<xref ref-type="bibr" rid="B5">Azzola et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B6">Beca-Carretero et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B8">Bellin and Rossi, 2024</xref>; <xref ref-type="bibr" rid="B41">Martin et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B47">Moraitis et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B64">Vassallo et&#xa0;al., 2018</xref>).</p></list-item>
</list>
<p>The usage of benthic habitat classification systems is fundamental (<xref ref-type="bibr" rid="B46">Montefalcone et&#xa0;al., 2021</xref>) to characterize and describe the habitats (<xref ref-type="bibr" rid="B54">Robinson and Levings, 1995</xref>). In particular, a classification scheme provides a structured framework for the description and standardization of the physical and biological conditions defining habitat classes (<xref ref-type="bibr" rid="B59">Strong et&#xa0;al., 2019</xref>).</p>
<p>Numerous Benthic Habitat Classification Schemes (BHCSs) have been developed with different goals around the world (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). Many of these schemes and lists are incompatible with each other, making it difficult to compare habitat types across studies and regions (<xref ref-type="bibr" rid="B32">Greene et&#xa0;al., 2008</xref>). Numerous scientific papers have reviewed existing classification systems for marine benthic habitats, discussed the revision process, and identified gaps (<xref ref-type="bibr" rid="B21">Diaz et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B26">Fraschetti et&#xa0;al., 2008</xref>; <xref ref-type="bibr" rid="B27">Galparsoro, 2012</xref>; <xref ref-type="bibr" rid="B45">Misiuk and Brown, 2024</xref>; <xref ref-type="bibr" rid="B46">Montefalcone et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B59">Strong et&#xa0;al., 2019</xref>).</p>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Non-exhaustive list of international, European and regional classification schemes.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center">Name</th>
<th valign="middle" align="center">Purpose</th>
<th valign="middle" align="center">Responsible party</th>
<th valign="middle" align="center">References</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="center">Nouveau manuel de bionomie benthique de la M&#xe9;diterran&#xe9;e</td>
<td valign="middle" align="center">A detailed account of Mediterranean communities often referred to as biocenoses, which are commonly used in the Mediterranean region to interpret distribution patterns of marine species and assemblages within a deterministic framework.</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B50">P&#xe9;r&#xe8;s, 1967</xref>; <xref ref-type="bibr" rid="B51">P&#xe9;r&#xe8;s and Picard, 1964</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">EUropean Nature Information System (EUNIS)</td>
<td valign="middle" align="center">Widely used for referencing and reporting habitat data in a consistent manner, supporting inventories, monitoring, assessments, and biodiversity indicators across Europe.</td>
<td valign="middle" align="center">European Environment Agency (EEA)</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B20">Davies et&#xa0;al., 2004</xref>; <xref ref-type="bibr" rid="B19">Davies and Moss, 1998</xref>)<break/><ext-link ext-link-type="uri" xlink:href="https://eunis.eea.europa.eu/habitats-code-browser-revised.jsp">https://eunis.eea.europa.eu/habitats-code-browser-revised.jsp</ext-link></td>
</tr>
<tr>
<td valign="middle" align="center">Barcelona Convention classification</td>
<td valign="middle" align="center">Developed to map and monitor marine habitats in the Mediterranean Sea in 1998 and revised in 2021 consistently with the criteria used for updating the EUropean Nature Information System (EUNIS).</td>
<td valign="middle" align="center">Barcelona Convention</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B46">Montefalcone et&#xa0;al., 2021</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">Potential Habitat Characterization Scheme (PHCS)</td>
<td valign="middle" align="center">To map marine benthic habitats in deep water using sensors data, video, photographs, and seafloor samples. It considers four spatial scales, and mainly it uses physical parameters and features to classify the seafloor.</td>
<td valign="middle" align="center">Moss Landing Marine Laboratories (MLML) Administered by San Jose State University</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B32">Greene et&#xa0;al., 2008</xref>, <xref ref-type="bibr" rid="B31">2005</xref>, <xref ref-type="bibr" rid="B30">1999</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">Classification of Sublittoral Habitats (CSH)</td>
<td valign="middle" align="center">Designed to classify marine sublittoral habitats in the American and Canadian regions of northeastern North America through the use of geophysical surveys, along with video and photographic transects, and sediment and biological sampling.</td>
<td valign="middle" align="center">United States Geological Survey and Natural Resources Canada</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B63">Valentine et&#xa0;al., 2005</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">Australian National Intertidal/Subtidal Benthic (NISB) Habitat Classification Scheme</td>
<td valign="middle" align="center">To categorize and map marine habitats in the intertidal and subtidal zones. Developed to provide a consistent method for identifying and classifying benthic habitats across Australia, aiding in marine management and conservation efforts.</td>
<td valign="middle" align="center">Australian Coastal Vulnerability Project</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B48">Mount et&#xa0;al., 2007</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">Coastal and Marine Habitat and Ecosystem Classification (CMHEC)</td>
<td valign="middle" align="center">To categorize and understand the diverse marine and coastal environments in New Zealand. It is a three-dimensional classification scheme, taking into account surface, water column and benthic features.</td>
<td valign="middle" align="center">Ministry of Fisheries (MFish) and Ministry of Fisheries and Department of Conservation (DOC)</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B44">Ministry of Fisheries and Department of Conservation, 2008</xref>; <xref ref-type="bibr" rid="B55">Rowden et&#xa0;al., 2018</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">Coastal and Marine Ecological Classification Standard (CMECS)</td>
<td valign="middle" align="center">Aimed at describing, classifying, organizing, and interpreting marine ecological data using a semi-hierarchical framework that incorporates various settings (aquatic and biogeographic) and components (biotic, water column, substrate, and geoforms).</td>
<td valign="middle" align="center">National Oceanic and Atmospheric Administration (NOAA)</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B40">Madden et&#xa0;al., 2009</xref>; <xref ref-type="bibr" rid="B58">Standards Working Group - Federal Geographic Data Committee, 2012</xref>)<break/><ext-link ext-link-type="uri" xlink:href="https://iocm.noaa.gov/standards/cmecs-home.html">https://iocm.noaa.gov/standards/cmecs-home.html</ext-link></td>
</tr>
<tr>
<td valign="middle" align="center">Hierarchical Framework of Marine Habitat Classification for Ecosystem-Based Management (HFMHC)</td>
<td valign="middle" align="center">A multi-scale hierarchical framework with emphasis on finer-scale habitat classification levels, offering conceptual schematics to guide habitat studies and inform management decisions.</td>
<td valign="middle" align="center">Graduate School of Oceanography, University of Rhode Island</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B33">Guarinello et&#xa0;al., 2010</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">El Inventario Espa&#xf1;ol de H&#xe1;bitats y Especies Marinos (IEHEM)</td>
<td valign="middle" align="center">To establish and hierarchically classify the around 890 habitats identified in the Spanish marine environment. IEHEM is part of another global inventory called the Spanish Inventory of Natural Heritage and Biodiversity regulated by Royal Decree 556/2011.</td>
<td valign="middle" align="center">Spanish Inventory of Natural Heritage and Biodiversity. established by Law 42/2007</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B61">Templado et&#xa0;al., 2012</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">HELCOM Underwater biotope and habitat classification system (HELCOM Hub)</td>
<td valign="middle" align="center">To define biotopes in the whole Baltic Sea. It is hierarchial and structured into six levels. Habitats are defined as the abiotic environment, while biotopes are defined as the abitotic environment coupled with the characteristic organism community.</td>
<td valign="middle" align="center">HELCOM Red List project</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B4">Avellan et&#xa0;al., 2013</xref>)<break/><ext-link ext-link-type="uri" xlink:href="http://www.helcom.fi/baltic-sea-trends/biodiversity/helcom-hub">http://www.helcom.fi/baltic-sea-trends/biodiversity/helcom-hub</ext-link></td>
</tr>
<tr>
<td valign="middle" align="center">Seamap Australia classification scheme</td>
<td valign="middle" align="center">To classify together living, non-living, and contextual components to define a seabed habitat.</td>
<td valign="middle" align="center">Australian National Data Service (ANDS) High Values Collection (HVC) program</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B11">Butler et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B39">Lucieer et&#xa0;al., 2017</xref>)<break/><ext-link ext-link-type="uri" xlink:href="https://seamapaustralia.org/resources/classification">https://seamapaustralia.org/resources/classification</ext-link></td>
</tr>
<tr>
<td valign="middle" align="center">CoralFISH hierarchical biotope classification scheme</td>
<td valign="middle" align="center">To reach a detailed taxonomic description of Cold-Water Corals (CWCs) in Europe.</td>
<td valign="middle" align="center">EU FP7 project CoralFISH</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B18">Davies et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B34">Guillaumont et&#xa0;al., 2016</xref>)</td>
</tr>
<tr>
<td valign="middle" align="center">IUCN Global Ecosystem Typology</td>
<td valign="middle" align="center">A comprehensive classification framework for Earth&#x2019;s ecosystems that integrates functional and compositional characteristics, with a focus on the IUCN Red List of Ecosystems.</td>
<td valign="middle" align="center">IUCN Regional Office for Mexico, Central America and the Caribbean (ORMACC) and the Commission on Ecosystem Management (CEM)</td>
<td valign="middle" align="center">(<xref ref-type="bibr" rid="B38">Keith et&#xa0;al., 2020</xref>)</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>This paper introduces the Coast to Deep Mapping (CoDeMap) benthic habitats classification scheme (BHCS) for the Mediterranean and Black Sea benthic habitats from the coast to the deep sea. The philosophy behind CoDeMap is to provide a practical and operative tool for users from different backgrounds who need to organize and interpret marine observational data, as well as to describe, classify, and map seafloors.</p>
</sec>
<sec id="s2">
<label>2</label>
<title>CoDeMap benthic habitats classification scheme</title>
<p>CoDeMap BHCS is inspired by already existing classification schemes (EUNIS, CMECS, Seamap Australia) and habitats lists (IEHEM, <xref ref-type="bibr" rid="B51">P&#xe9;r&#xe8;s and Picard (1964)</xref>, Annex II of the Habitats Directive, <xref ref-type="bibr" rid="B61">Templado et&#xa0;al. (2012)</xref>) with a focus on the commonly underrepresented mesophotic and deep-sea environments. The aim during development of the CoDeMap was to create a classification scheme:</p>
<list list-type="simple">
<list-item>
<p>- Scientifically-based but easily applicable, with separated abiotic and biotic components defining the benthic habitat to minimize the uncertainties and biases introduced with the subjective interpretation;</p></list-item>
<list-item>
<p>- Hierarchical, its components are organized in subcomponents and sublevels able to catch the complexity of seafloor according to the availability and quality level of spatial data;</p></list-item>
<list-item>
<p>- Multiscale, user can capture the most relevant scale-dependent patterns and the high complexity and spatial heterogeneity of the seafloor, encompassing both abiotic and biotic characteristics;</p></list-item>
<list-item>
<p>- Multipurpose, CoDeMap is compatible with all mapping techniques. Users can map according to (i) typology, availability, and quality of the spatial data, (ii) the user expertise, (iii) the target (abiotic maps, single biota maps, community maps, benthoscape maps), and (iv) the purpose of the spatial representation (e.g. scientific papers, monitoring activities, inventory, prediction models, legislation background, habitat-based management measures);</p></list-item>
<list-item>
<p>- Flexible, CoDeMap is primarily designed for the Mediterranean and Black Sea, but it could be easily applied to any marine situation worldwide through the adaptations of its codes; the ability to combine classes allows for the description of habitat mosaics, enabling more accurate representation of seabed conditions that do not fit neatly into predefined classes;</p></list-item>
<list-item>
<p>- Dynamic and public, CoDeMap is publicly available (<ext-link ext-link-type="uri" xlink:href="https://codemap.my.canva.site/about">https://codemap.my.canva.site/about</ext-link>) including versioning and a form for the contribute implementing of the scheme. Indeed, it provides a baseline suitable to be constantly updated addressing increase of knowledge and predictable future changes of marine ecosystems.</p></list-item>
</list>
<sec id="s2_1">
<label>2.1</label>
<title>Components, subcomponent, levels and classes</title>
<p>The CoDeMap scheme is organized into three main components: 1) Morphology, 2) Substrate, and 3) Biology (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>).</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>The three components of the CoDeMap benthic habitat classification scheme (Morphology, Substrate and Biology).</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g001.tif">
<alt-text content-type="machine-generated">Flowchart illustrating the components of a benthic habitat map. Step 1: Morphological component, depicted with physiography, geoforms, and bedforms, in orange. Step 2: Substrate component, represented in yellow. Step 3: Biological component, shown in green. These steps lead to the resulting benthic habitat map, highlighted in blue.</alt-text>
</graphic></fig>
<p>Internally, the main components are systemized hierarchically, with a series of subcomponents, levels, and classes (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). Seafloor morphology, type of substrate, and distribution of individual species or communities can be mapped separately and then merged into a single map of benthic habitats by using GIS software.</p>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p>Structure of the CoDeMap BHCS showing the three components (red squares), the sub-components (yellow squares), and the levels (green circles). The levels contain the classes defined by a code and a label.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g002.tif">
<alt-text content-type="machine-generated">Diagram showing the CoDeMap framework with components: Morphology, Substrate, and Biology. Morphology has sub-components Physiography, Geoforms, Bedforms, leading to levels PL1, GL1-3, BFL1. Substrate leads to levels SL1-3. Biology leads to levels BL1-3. Classes include examples like Continental slope, Canyon, Rock substrate, and Coral reef.</alt-text>
</graphic></fig>
<p>More specifically, the CoDeMap morphology components consist of subcomponents organized in descending order of size and level of detail 1) Physiography, 2) Geoforms, and 3) Bedforms. Within each subcomponent, levels are used to classify features from a broader scale (Level 1, L1) to small-scale features (L2, L3). In particular, the Physiography sub-component contains one level (PL1), while the Geoforms sub-component contains three levels (GL1, GL2, and GL3), and the Bedforms sub-component contains&#xa0;one level (BFL1). The Substrate and Biology components include three levels each, SL1, SL2, SL3 and BL1, BL2, BL3 respectively (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>).</p>
<p>The sum of the three components (Morphology, Substrate and Biology) returns a benthic habitat map where each class is uniquely and unequivocally represented. The maximum number of levels describing a habitat class is 11, but not all levels must perforce contain information.</p>
<p>Each class of the scheme is identifiable by a univocal alphanumeric code and a label (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2</bold></xref>). Both the complexity of the code and the detail expressed with the label of the features increase from L1 to L3. Within L3 of substrate and biology, classes can be more specific (e.g. B020302 &#x2013; <italic>Codium adherens</italic>) or more generic (e.g. B020301- Green algae) in bold in the scheme. To limit the proliferation and redundancy of classes in the scheme, codes can be combined to describe situations characterized by multiple classes, ordered by prevalence. The combination is permitted if more classes coexist and there is a representativeness of at least 25%. For example, an area characterized by coralligenous (spatial coverage=75%) and <italic>Posidonia oceanica</italic> (spatial coverage=25%) can be described as B0907+B040403. Conversely, a <italic>Posidonia oceanica</italic> meadow (coverage=75%) with interspersed coralligenous outcrops (coverage=25%) can be codified as B040403+B0907.</p>
<p>The legend can be customized to include only codes, only labels, or a combination of both. Additionally, users can select which levels to display according to the complexity and purpose of the representation (see paragraph 3).</p>
<sec id="s2_1_1">
<label>2.1.1</label>
<title>Morphological component</title>
<p>Within the Morphology component, the Physiography sub-component includes only level PL1, which comprises the different constituents of the continental margin (<italic>i.e</italic>., moving from shallow to deep areas, coast, shelf, continental slope, basin plain, etc). The Geoform sub-component is divided into three levels (GL1, GL2, GL3). GL1 concerns environments and large-scale morphological features (e.g. beach, submarine canyon, leveed channels). GL2 refers to medium-scale morphologies and/or sub-environments (e.g. foreshore, canyon flank), while GL3 considers small-scale morphologies (e.g. shoreface bar, intra-canyon plunge pool, intra-lobe channel). For example, in the CoDeMap scheme a terrace on a canyon flank is coded G090201, (G09 - Canyon, G0902 - Flank, G090201 - Terrace). The Bedform sub-component consists of 17 features at a single level (encoded BF01, BF02, BF03, etc.). Therefore, a morphological feature can be described hierarchically using a complete code consisting of the union of sub-components. If along the continental slope (P05) a canyon (G09), whose flank (G0902) is marked by several incisions (BF16) the final code will result in: P02G0902BF16. In order to gather and organize all these classes together, we considered works like <xref ref-type="bibr" rid="B3">Ashley (1990)</xref>; <xref ref-type="bibr" rid="B22">Dove et&#xa0;al. (2020)</xref>; <xref ref-type="bibr" rid="B36">Harris et&#xa0;al. (2014)</xref>, and <xref ref-type="bibr" rid="B43">Micallef et&#xa0;al. (2018)</xref>.</p>
</sec>
<sec id="s2_1_2">
<label>2.1.2</label>
<title>Substrate component</title>
<p>The Substrate component classifies seabed nature and consists of three levels (SL1, SL2, SL3). SL1 distinguishes between consolidated (i.e. hard substrate), unconsolidated (i.e. soft substrate), and semi-consolidated substrate (i.e. various stages of lithification). SL2 provides information about the type of seabed (e.g. rocky substrate, firmground, biogenic unconsolidated substrate), similarly to the CMECS. SL3 considers the grain size (e.g. gravel, sand, mud) according to <xref ref-type="bibr" rid="B66">Wentworth (1922)</xref>, and the type of sediment (e.g. cohesive mud, bioclastic sand, coral rubble). Therefore, an area characterized by blocks and boulders is coded as Consolidated substrate (S01), Rocky substrate (S0101), and Block and boulder (S010102).</p>
</sec>
<sec id="s2_1_3">
<label>2.1.3</label>
<title>Biological component</title>
<p>The Biology component consists of three levels (BL1, BL2, BL3). The coarsest level (BL1) considers different morpho-functional groups representing the seascape, (e.g., turf, forest, bioconstruction). BL2 specifies broad taxonomic groups represented in BL1. While, BL3 includes the highest possible taxonomic level, genus or species, (e.g., <italic>Callogorgia verticillata</italic>), or morpho-functional groups of species (e.g., red algae and massive sponges). BL3 has been conceived to allow experts and non-expert users to document more detailed biodiversity information. For example, the code B090737 indicates a Bioconstruction (B09) made by Coralligenous (B0907) characterized by Massive sponges (B090737).</p>
<p>Considering a mosaic of habitats characterized by the co-occurrence in high number of <italic>Madrepora oculata</italic> (B090210) and <italic>Poecillastra compressa</italic> (B070222) (coverage=60% and 40%, respectively), it should be categorized as &#x201c;<italic>M. oculata&#x201d;</italic> + &#x201c;<italic>P. compressa&#x201d;</italic> (B090210+B070222). If the user cannot (or is not able to) recognize a single species or several taxa typifying the area, it is also possible to mix different levels of the component. Following the previous example, a coral reef made by <italic>M. oculata</italic> (coverage=60%) + and a ground dominated by massive sponge where it is not possible to recognize the dominant species (coverage=40%) will be coded as B090210+B0702. The order of the two classes is related to their relative abundance. It is possible to reverse the codes if the coverage is different: B0702+B090210 identified an area characterized by a massive sponge ground (coverage=60%) and a coral reef built by <italic>M</italic>. <italic>oculata</italic> (coverage=40%).</p>
<p>The possibility to mix classes from different components permits the user to classify each item with a unique code in CoDeMap. Meaning what the <italic>Posidonia oceanica</italic> is identified by the code B040203, which can be associated with different substrate types (e.g. <italic>P. oceanica</italic> on sand is classified as S030301B040203 and <italic>P. oceanica</italic> on <italic>matte</italic> is classified as S030105B040203).</p>
<p>Several classification schemes and lists have been considered to compile the biological components, among others: EUNIS, IEHEM, Annex II of the Habitat Directive 92/43/EEC, Classification of benthic marine habitat types for the mediterranean region (SPA/RAC, 2006) and, IUCN Red List of Threatened Species.</p>
</sec>
</sec>
</sec>
<sec id="s3">
<label>3</label>
<title>GIS applications</title>
<p>This section describes four applications of the CoDeMap BHCS for different mapping scenarios, which are characterized by different scales, knowledge backgrounds and purposes. In the first application, the tool is used to create a large-scale map of benthic habitats for the Southern Adriatic (Mediterranean Sea) based on indirect and inhomogeneous geophysical data. The second example uses CoDeMap to describe the Tricase Canyon (Adriatic Sea), considering all three components of the scheme. The third application, CoDeMap is used to map the seafloor of the Dohrn Canyon (Tyrrhenian Sea) that has been surveyed by a Remotely Operated Vehicle (ROV). Finally, the fourth case study is a comparison between the CoDeMap and EUNIS classification schemes in the continental shelf along the Apulian coast (South Adriatic Sea).</p>
<sec id="s3_1">
<label>3.1</label>
<title>South Adriatic continental margin</title>
<p>The South Adriatic Sea has been investigated by the CNR-ISMAR throughout the last 20 years by the acquisition of a large amount of geophysical data (multibeam and seismic), seabed samples (grab samples, box cores), and video from ROV. The interpretation of these data provided the basis to produce a geomorphological map of the South Adriatic continental margin (<xref ref-type="bibr" rid="B12">Campiani et al., 2024</xref>), and a benthic habitat map published in <xref ref-type="bibr" rid="B52">Prampolini et&#xa0;al. (2021)</xref>. In this application, we have classified these two products using the CoDeMap BHCS producing several maps representing the morphology, the substrate, the biology, and the benthic habitat map of the basin.</p>
<p><xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3A</bold></xref> represents the Morphology component classified according to the Physiography sub-component. In <xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3B</bold></xref>, the morphologic classification includes the level of Physiography (PL1) and the three levels of Geoforms (GL1, GL2, GL3), increasing the detail and the complexity of the seascape. The complete South Adriatic morphology is charted in <xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4</bold></xref>, where all the sub-components (Physiography, Geoforms and Bedforms) were used to build the map. These three representations of the South Adriatic morphology enhance the increase in scale, detail and complexity of the depicted seascape, by applying different levels of the CoDeMap BHCS and consequently, changing the information represented on the map.</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p>Morphology of the South Adriatic continental margin classified according to CoDeMap BHCS: <bold>(A)</bold> Physiography sub-component, <bold>(B)</bold> Physiography and Geoforms sub-components. Background: EMODnet Bathymetry World Base Layer version 1.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g003.tif">
<alt-text content-type="machine-generated">Two maps of the South Adriatic Sea display physiography and geoforms with legends. Map A shows broad categories: continental shelf (green), continental slope (blue), and basin plain (purple). Map B provides detailed geoform distinctions, including features like sand volcanoes, channels, and erosive remnants. Both maps are oriented north with Italy discernible to the west. The legends define color codes for each landform type.</alt-text>
</graphic></fig>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Morphology of the South Adriatic continental margin according to CoDeMap BHCS: full classification of the Morphology component using all the three sub-components: Physiography, Geoforms and Bedforms. For each element of the legend, the full code is shown coupled with the label of the most detailed class of the CoDeMap BHCS. Background: EMODnet Bathymetry World Base Layer version 1.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g004.tif">
<alt-text content-type="machine-generated">Map of the South Adriatic Sea region, illustrating physiography and geoforms. Various colors and patterns represent different features like continental shelves, slopes, and basin plains. Key geographical locations include Italy and Albania. A legend identifies features such as sediment wave areas, canyons, and depositional zones. A scale bar indicates distances.</alt-text>
</graphic></fig>
<p>The South Adriatic continental margin has been classified according to information on substrate and biology at different levels of detail. The map in <xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref> derives from the interpretation of seabed reflectivity and samples; the latter permitted to specify substrate texture and biological communities living on the seafloor. The substrate is described by all levels of the CoDeMap Substrate component and represented by using distinct colors range and tones according to the texture. The Biology component is depicted through a halftone screen superimposed on the substrate.</p>
<fig id="f5" position="float">
<label>Figure&#xa0;5</label>
<caption>
<p>Substrate and Biology components of the South Adriatic continental margin classified according to CoDeMap BHCS (modified from <xref ref-type="bibr" rid="B52">Prampolini et&#xa0;al., 2021</xref>). Background: EMODnet Bathymetry World Base Layer version 1.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g005.tif">
<alt-text content-type="machine-generated">Map of the Adriatic Sea region showing various habitats marked by colors and patterns. Insets focus on detailed areas. A color-coded legend identifies 29 habitats such as coral reefs, mud, and bioclastic sand. These are labeled with codes for substrates and biological features. A scale bar indicates distances, and the map includes geographical coordinates. The Adriatic Sea is labeled, with clear sectioning of terrestrial and marine areas.</alt-text>
</graphic></fig>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Tricase Canyon</title>
<p>The Tricase Canyon is a submarine feature that cuts through the Apulian continental slope on the western side of the Ionian Sea (Mediterranean Sea). Morphological and substrate components were mapped by interpreting the Digital Elevation Model (DEM) and classifying seabed acoustic reflectivity using Remote Sensor Object-Based Image Analysis (RSOBIA). A ground-truthing activity involving seabed samples and ROV images validated the results of the automatic classification and helped analyze the biological component. In fact, the deeper areas of the canyon host white corals such as <italic>Madrepora oculata</italic>, <italic>Desmophyllum dianthus</italic>, and <italic>Desmophyllum pertusum</italic> (=<italic>Lophelia pertusa</italic>). Corals have been mainly observed on the top of blocks interpreted as the result of several mass-transport deposits (<xref ref-type="bibr" rid="B53">Prampolini et&#xa0;al., 2020</xref>). <xref ref-type="fig" rid="f6"><bold>Figure&#xa0;6</bold></xref> shows the benthic habitat map of the Tricase Canyon described using the CoDeMap BHCS.</p>
<fig id="f6" position="float">
<label>Figure&#xa0;6</label>
<caption>
<p>Benthic habitat map of the Tricase Canyon classified according to CoDeMap (codes explanation is given in <xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). Colors represent the morphologies of the canyon (yellow tones for the continental shelf, orange-green tones for the continental slope and blue tones for the basin plain); the pattern of the polygons corresponds to a specific substrate that is red when also the biological component is present. Background: EMODnet Bathymetry World Base Layer version 1.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g006.tif">
<alt-text content-type="machine-generated">Map depicting the seabed morphology, substrate, and biology of an area off the coast of Italy. Different colored patterns represent various geological and biological classifications, as detailed in the legend. The inset illustrates the location within the Mediterranean region. The map includes a scale bar and orientation compass, showing latitude and longitude coordinates.</alt-text>
</graphic></fig>
<p>For the Tricase Canyon case, we decided to use all 11 levels in the legend displaying just the code, integrating all three components into a single seafloor representation. <xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref> contains the labels for the various classes.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Description of the codes used in the legend of <xref ref-type="fig" rid="f6"><bold>Figure&#xa0;6</bold></xref>.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="left">Component</th>
<th valign="middle" align="left">Code</th>
<th valign="middle" align="left">Label</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P02</td>
<td valign="middle" align="left">Continental shelf</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P02G0706</td>
<td valign="middle" align="left">Continental shelf - Cold-seep feature - Sand volcano</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05</td>
<td valign="middle" align="left">Continental slope</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G0901</td>
<td valign="middle" align="left">Continental slope - Canyon - Head</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G0902</td>
<td valign="middle" align="left">Continental slope - Canyon - Flank</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G0902BF16</td>
<td valign="middle" align="left">Continental slope - Canyon - Flank - Incision</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G0903</td>
<td valign="middle" align="left">Continental slope - Canyon - Floor</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G1001</td>
<td valign="middle" align="left">Continental slope - Channel - Thalweg</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G1007</td>
<td valign="middle" align="left">Continental slope - Channel - Levee</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G12</td>
<td valign="middle" align="left">Continental slope - Mass transport deposit</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P05G120301</td>
<td valign="middle" align="left">Continental slope - Mass transport deposit - Depositional zone - Slide block</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P07</td>
<td valign="middle" align="left">Basin plain</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P07G0904</td>
<td valign="middle" align="left">Basin plain - Canyon - Thalweg</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P07G1203</td>
<td valign="middle" align="left">Basin plain - Mass transport deposit - Depositional zone -</td>
</tr>
<tr>
<td valign="middle" align="left">Morphology</td>
<td valign="middle" align="left">P07G120301</td>
<td valign="middle" align="left">Basin plain - Mass transport deposit - Depositional zone - Slide block</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S010101</td>
<td valign="middle" align="left">Consolidate substrate - Rocky substrate - Bedrock</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S010102</td>
<td valign="middle" align="left">Consolidate substrate - Rocky substrate - Block and boulder</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S010104</td>
<td valign="middle" align="left">Consolidate substrate - Rocky substrate - Lithified sediment</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S030306</td>
<td valign="middle" align="left">Unconsolidated substrate - Fine unconsolidated substrate - Bioclastic muddy sand</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S030309</td>
<td valign="middle" align="left">Unconsolidated substrate - Fine unconsolidated substrate - Sandy mud</td>
</tr>
<tr>
<td valign="middle" align="left">Substrate</td>
<td valign="middle" align="left">S030313</td>
<td valign="middle" align="left">Unconsolidated substrate - Fine unconsolidated substrate - Mud</td>
</tr>
<tr>
<td valign="middle" align="left">Biology</td>
<td valign="middle" align="left">B090208</td>
<td valign="middle" align="left">Bioconstruction &#x2013; Coral reef &#x2013; <italic>Desmophyllum pertusum</italic></td>
</tr>
<tr>
<td valign="middle" align="left">Biology</td>
<td valign="middle" align="left">B090209</td>
<td valign="middle" align="left">Bioconstruction &#x2013; Coral reef &#x2013; <italic>Desmophyllum dianthus</italic></td>
</tr>
<tr>
<td valign="middle" align="left">Biology</td>
<td valign="middle" align="left">B090210</td>
<td valign="middle" align="left">Bioconstruction &#x2013; Coral reef &#x2013; <italic>Madrepora oculata</italic></td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>ROV transect in the Dohrn Canyon</title>
<p>The Dohrn Canyon is in the center of the Gulf of Naples, a submarine canyon of ecological, functional and oceanographic interest since featured by important upwelling currents affecting more coastal waters. It hosts deep bioconstructions, specifically cold-water corals and oysters. Available information documents the presence of living specimens of the scleractinians <italic>M. oculata</italic>, <italic>D. pertusum</italic>, and <italic>D. dianthus</italic>. These communities are also associated with large bivalves such as <italic>Neopycnodonte zibrowii</italic> and <italic>Acesta excavata</italic>. This coexistence of deep corals and large bivalves represents a unique biotope for the Mediterranean Sea, threatened by severe anthropogenic threats (<xref ref-type="bibr" rid="B60">Taviani et&#xa0;al., 2019</xref>).</p>
<p>In this application of the CoDeMap BHCS, we described the ROV transect coded &#x201c;ANOMCITY_ROV01&#x201d;, conducted during the CNR oceanographic cruise ANOMCITY 2016 aimed to characterize and map the bioconstructions populating the Dohrn Canyon (<xref ref-type="bibr" rid="B49">Oliveri et&#xa0;al., 2016</xref>). The transect develops along the flank of the northern branch of the canyon following a South-North direction, revealing the coexistence of cold-water corals and deep oysters (<xref ref-type="fig" rid="f7"><bold>Figure&#xa0;7</bold></xref>).</p>
<fig id="f7" position="float">
<label>Figure&#xa0;7</label>
<caption>
<p><bold>(A, B)</bold> show the location of the ROV transect ANOMCITY_ROV01 in the Mediterranean Sea and in the Dohrn Canyon, respectively. In <bold>(C)</bold>, the seafloor is described using the three components of the CoDeMap BHCS. <bold>(D)</bold> shows a zoom in the rocky part of the transect hosting cold-water corals (B0902 - Bioconstructions, Coral reef) and deep oyster (B0903 - Bioconstruction, Oyster s.l. reef). Point size refers to the abundance of specimens.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g007.tif">
<alt-text content-type="machine-generated">Map series showing geological and biological data of the Mediterranean region. Map A displays a broader view of Europe and the Mediterranean Sea, indicating specific study locations. Map B zooms into a selected area, highlighting undersea topography. Maps C and D provide detailed bathymetric data with contour lines and tracks of ROV (remotely operated vehicle) dives, marked in various colors representing specific routes and specimen abundances. Blue, yellow, and green circles indicate different species and their numbers, according to the legend detailing specimen abundance. Locations are marked with north arrows and scale bars.</alt-text>
</graphic></fig>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Continental shelf along the Apulian coast</title>
<p>From 2000 to 2024 several research projects and scientific papers focused on the area in the South Adriatic Sea that runs along the Apulian coast and continental shelf (Italy). In this case study, we considered the seabed stretching from Mola di Bari to Fasano municipalities hosting <italic>Posidonia oceanica</italic> meadows, coralligenous bioconstructions and coral reefs in shallow waters, as well as deep oyster reefs in the continental shelf at approximately at 100 meters water depth. The seabed substrate ranges from hardgrounds to bioclastic coarse and fine sediments (from gravelly sands to sandy muds), while the continental shelf is characterized by flat surfaces, megaripple fields, comet marks, and erosional remnants. We integrated the maps produced through the years from the coast to the deep-sea into a comprehensive map intended to support conservation initiatives, such as the establishment of new Natura 2000 sites (<xref ref-type="bibr" rid="B29">Grande et&#xa0;al., 2024</xref>). The main challenge lies in homogenizing maps derived from multiple sources, produced at different scales and using different devices (see <xref ref-type="supplementary-material" rid="SF1"><bold>Supplementary Figure S1</bold></xref> in the <xref ref-type="supplementary-material" rid="ST1"><bold>Supplementary Materials</bold></xref> for the original data).</p>
<p><xref ref-type="fig" rid="f8"><bold>Figure&#xa0;8</bold></xref> compares the CoDeMap and EUNIS classification schemes in the production of benthic habitat maps for Apulian coastal waters and continental shelf. Both frameworks ensure a consistent representation of the study area, even in contexts where data availability is limited. Within EUNIS, these areas can be classified at Level 1 as &#x201c;Marine benthic habitats,&#x201d; whereas CoDeMap adds further detail by incorporating physiographic features such as the &#x201c;Continental shelf.&#x201d; The key distinction between the two approaches lies in the reliance on biozones in EUNIS versus geomorphological classes in CoDeMap. In <xref ref-type="fig" rid="f8"><bold>Figure&#xa0;8A</bold></xref>, geomorphological features are clearly delineated, with biological data embedded within their abiotic setting. For instance, the oyster reef offshore Monopoli is shown to coincide with an erosive remnant area, offering valuable insights into the reef&#x2019;s formation. Conversely, EUNIS highlights the spatial distribution of biozones: <xref ref-type="fig" rid="f8"><bold>Figure&#xa0;8B</bold></xref> clearly illustrates the extent of habitats across the infralittoral and circalittoral zones.</p>
<fig id="f8" position="float">
<label>Figure&#xa0;8</label>
<caption>
<p>Comparison between the CoDeMap and EUNIS classification schemes applied on the South Adriatic Sea continental shelf along the Apulian coast (Italy). <bold>(A)</bold> represents the benthic habitats classified according to the CoDeMap BHCS, and <bold>(B)</bold> the same habitats classified according to the EUNIS classification scheme.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-12-1663369-g008.tif">
<alt-text content-type="machine-generated">Two marine benthic habitat maps showing different classification systems over the same coastal area. The top map uses the CoDeMap classification, with labels indicating features like continental shelf, coral reefs, and various sediment types. The bottom map uses the EUNIS classification, highlighting habitats such as marine benthic habitats, coralligenous biocenosis, and Mediterranean substrate variations. Each map features a legend with code-label pairs for reference. The maps include detailed color-coded sections representing various geophysical and biological features, and both include a scale bar indicating a distance of 10 kilometers.</alt-text>
</graphic></fig>
<p>In terms of completeness, CoDeMap generally provides a more detailed account of the original data. This is exemplified by the mesophotic coral reef described by <xref ref-type="bibr" rid="B16">Corriero et&#xa0;al. (2019)</xref>, classified under CoDeMap as a coral reef dominated by <italic>Phyllangia americana mouchezii</italic> and <italic>Polycyathus muellerae</italic>, whereas EUNIS categorizes it as &#x201c;MC2 &#x2013; Mediterranean circalittoral biogenic habitat.&#x201d; In this case, the absence of specific EUNIS classes necessitates classification at Level 3, resulting in a coarser description. By contrast, CoDeMap captures the reef&#x2019;s character more precisely, including the identification of its dominant species.</p>
<p>When harmonizing maps from different sources, however,&#xa0;some information may be lost, potentially leading to misclassification or the use of categories that do not fully match the context. This is illustrated in <xref ref-type="fig" rid="f8"><bold>Figure&#xa0;8</bold></xref> with the mapping of <italic>Posidonia oceanica</italic> from the &#x201c;Inventory and Cartography of Posidonia Meadows&#x201d; (POR 2000&#x2013;2006). The dataset shows a mosaic o<italic>f Posidonia oceanica</italic> and matte within the circalittoral zone. Under the EUNIS scheme, the area can only be mapped as MB252 &#x201c;Biocenosis of <italic>Posidonia oceanica</italic>&#x201d; (<xref ref-type="fig" rid="f8"><bold>Figure&#xa0;8B</bold></xref>), which omits details on the presence of <italic>matte</italic>, since the available data are insufficient to classify the habitat at Level 5 (e.g., MB2523 &#x201c;Facies of dead mattes of <italic>Posidonia oceanica</italic> without much epiflora&#x201d;). CoDeMap, by contrast, allows classes to be combined, thus retaining the full complexity of the original dataset.</p>
<p>Finally, CoDeMap consistently provides detailed information on substrate composition (e.g., gravelly and muddy sands or sandy muds across the continental shelf), whereas EUNIS categories do not always capture substrate variability exhaustively&#x2014;for example, &#x201c;MC45 Mediterranean circalittoral mixed sediment&#x201d;.</p>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>Over the past few decades, numerous classification systems have been developed, resulting in a variety of schemes and lists that are used for habitat description and monitoring. Many of these are incompatible among each other, making it difficult to compare habitat types across studies and regions (<xref ref-type="bibr" rid="B25">Fraschetti et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B32">Greene et&#xa0;al., 2008</xref>). The selection of the classification system to map benthic habitats is dependent on national preferences, established practices, and user expertise. Classifying natural continuities and environmental gradients into discrete and meaningful categories is a challenging endeavor, as it imposes constraints and limitations on the natural variability of ecological communities. Consequently, multiple BHCSs exist, differing in (i) purpose; (ii) environmental and ecological scope; (iii) spatial scale; (iv) thematic resolution; (v) structure; and (vi) compatibility for habitat mapping. Variations in these properties can significantly influence the presence and representation of marine habitat distributions (<xref ref-type="bibr" rid="B59">Strong et&#xa0;al., 2019</xref>).</p>
<p>CoDeMap is hierarchical and multiscale, adaptable to data availability and to the scale of work, and it is easy to manage and apply in a GIS environment. By using the CoDeMap BHCS, it is possible to map the different components of the benthic habitats separately and at different scales as shown in the application &#x201c;South Adriatic continental margin&#x201d; (<xref ref-type="fig" rid="f3"><bold>Figures&#xa0;3</bold></xref>&#x2013;<xref ref-type="fig" rid="f5"><bold>5</bold></xref>). This ensures the production of continuous maps for one or more components, regardless of the quality or quantity of the available data. Components can be combined to produce a benthic habitat map to get a full picture of a marine seafloor as demonstrated in the &#x201c;Tricase Canyon&#x201d; (<xref ref-type="fig" rid="f6"><bold>Figure&#xa0;6</bold></xref>) or according to detailed levels (<xref ref-type="fig" rid="f7"><bold>Figure&#xa0;7</bold></xref>). It is important to note that the resolution achievable within each component and level depends on the means and techniques employed for habitat mapping. For instance, the use of multibeam echosounder (MBES) data generally allows reliable classification of seafloor morphology down to the Geoform and Bedform levels (GL1&#x2013;3, BFL1), whereas sediment samples and ground-truthing techniques are essential to resolve Substrate levels (SL2&#x2013;3). Similarly, biological samples or seabed pictures are typically required to define the Biology component (BL1&#x2013;3), with biological samples necessary to reach the most detailed level of biology (BL3). Thus, the scheme provides a flexible framework where the depth of classification is directly related to the type and resolution of the available data.</p>
<p>Habitat mapping is a multidisciplinary endeavor, requiring collaboration among geologists, biologists, and other specialists. However, the modular structure of the CoDeMap scheme&#x2014;organized into separate components&#x2014;enables users to apply it according to their own expertise. For instance, if a geologist is unable to classify the biological component, the output will consist of a map of seabed morphology and substrate, which can later be complemented with biological information once a collaboration with biologists is established. Conversely, a biologist can map the biological component, with geological features subsequently added in partnership with geologists. This flexibility is not possible with classification schemes based on predefined combinations of components: a geologist would struggle to select among classes that share the same morphology and substrate but differ in biological communities, while a biologist would face difficulties distinguishing between classes defined by the same community but varying morphological or substrate characteristics.</p>
<p>In addition to scale and data availability, another driver in the application of CoDeMap can be the user purpose. If the focus is on geology, for instance, the morphology of the benthic habitat map can be the primary information displayed resulting in a continuous, colorful basemap, as illustrated in <xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4</bold></xref>. If the emphasis is on biology, substrate can be used as background element, then highlighting the biological elements layered on top (<xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref>). This is a great advantage because it allows the benthic habitat map representation to be updated or changed to suit the needs of any given project by simply accentuating a particular component above the others or by choosing the levels of interest. For example, in the application named &#x201c;ROV transect in the Dohrn Canyon&#x201d; (<xref ref-type="fig" rid="f7"><bold>Figure&#xa0;7</bold></xref>), the goal was to describe the seafloor characteristics and the biological community along an ROV transect for monitoring and conservation purposes. Finally, the ability to build purpose-driven maps makes CoDeMap a valuable tool in decision-making processes for users with varying levels of expertise and diverse backgrounds. Thanks to its immediacy and simplicity in conveying information, policy makers can also take advantage of CoDeMap: it enables them to clearly represent the messages and priorities they wish to communicate, thus facilitating understanding and the sharing of strategic decisions.</p>
<p>Such flexibility makes CoDeMap a user-friendly tool, enabling the classification of seafloor at various degrees. The applications of the CoDeMap BHCS highlight the scheme&#x2019;s versatility regarding spatial scale and code customization to suit the objectives of the representation. It is conceived as an evolving system that can be continuously enriched with new classes also to accommodate changes in future marine environments based on scientific community feedback (<xref ref-type="bibr" rid="B1">Albano et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B15">Coll et&#xa0;al., 2010</xref>; <xref ref-type="bibr" rid="B62">Thi&#xe9;bault and Moatti, 2016</xref>). In such a perspective, contributors can utilize a dedicated website (<ext-link ext-link-type="uri" xlink:href="https://codemap.my.canva.site/about">https://codemap.my.canva.site/about</ext-link>), where the latest version of the scheme is always accessible, and suggestions can be submitted.</p>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>In this work, we present CoDeMap, a classification scheme tailored for Mediterranean and Black Sea benthic habitats, ranging from coastal areas to the deep sea. CoDeMap offers a flexible framework for classifying marine benthic habitats suitable for GIS applications. It is rooted in scientific principles yet adaptable for various contexts, including citizen science, scientific research, and decision making. This study details the components, subcomponents, levels, and classes of CoDeMap, along with four diverse use cases that demonstrate the scheme&#x2019;s versatility in a range of scenarios (from simple assessments to highly detailed representations), according to scale, data availability, and individual expertise and objectives. CoDeMap will undergo continuous updates to reflect the dynamic nature of benthic habitats and marine ecosystem changes.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The maps used for GIS applications are freely accessible through the Geoportal for Marine Biodiversity in Italy, reachable via the Biodiversity Gateway (<uri xlink:href="https://www.biodiversitygateway.it">https://www.biodiversitygateway.it</uri>). The CoDeMap benthic classification scheme is available to download as an Excel file in the <xref ref-type="supplementary-material" rid="SM1"><bold>Supplementary Materials</bold></xref>. Different versions and an online contribution form are accessible at the following link: <uri xlink:href="https://codemap.my.canva.site/about">https://codemap.my.canva.site/about</uri>.</p></sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>VG: Conceptualization, Methodology, Software, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Validation. LA: Conceptualization, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Methodology, Validation. MP: Software, Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Validation. GCa: Writing &#x2013; original draft, Writing &#x2013; review &amp; editing, Validation. GV: Writing &#x2013; original draft, Validation. SF: Conceptualization, Funding acquisition, Supervision, Writing &#x2013; review &amp; editing. DBa: Validation, Writing &#x2013; review &amp; editing. DBe: Validation, Writing &#x2013; review &amp; editing. VB: Validation, Writing &#x2013; review &amp; editing. FC: Validation, Writing &#x2013; review &amp; editing. GCh: Validation, Writing &#x2013; review &amp; editing. AF: Validation, Writing &#x2013; review &amp; editing. BG: Validation, Writing &#x2013; review &amp; editing. FM: Validation, Writing &#x2013; review &amp; editing. MS: Validation, Writing &#x2013; review &amp; editing. AS: Validation, Writing &#x2013; review &amp; editing. PS: Validation, Writing &#x2013; review &amp; editing. VT: Validation, Writing &#x2013; review &amp; editing. MT: Conceptualization, Writing &#x2013; review &amp; editing, Validation. FF: Conceptualization, Funding acquisition, Methodology, Supervision, Writing &#x2013; review &amp; editing.</p></sec>
<ack>
<title>Acknowledgments</title>
<p>The Authors are grateful to the CoCoNet Consortium that participated in the first draft of the classification scheme. This is ISMAR-CNR, Bologna, scientific contribution n. 2091.</p>
</ack>
<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>
<p>The author(s) declared that they were an editorial board member of Frontiers, at the time of submission. This had no impact on the peer review process and the final decision.</p></sec>
<sec id="s10" sec-type="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p></sec>
<sec id="s11" 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="s12" sec-type="supplementary-material">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/fmars.2025.1663369/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/fmars.2025.1663369/full#supplementary-material</ext-link></p>
<supplementary-material xlink:href="Table1.xlsx" id="ST1" mimetype="application/vnd.openxmlformats-officedocument.spreadsheetml.sheet"><label>Supplementary Table&#xa0;S1</label>
<caption>
<p>CoDeMap benthic habitat classification scheme &#x2013; version 1.0..</p>
</caption></supplementary-material>
<supplementary-material xlink:href="Image1.jpg" id="SF1" mimetype="image/jpeg"><label>Supplementary Figure&#xa0;S1</label>
<caption>
<p>Data sources used for the case study &#x201c;Comparison between CoDeMap and EUNIS classification schemes&#x201d; paragraph 3.4. In red, the distribution of bioconstructions mapped in 2012 as part of the BIOMAP project (<ext-link ext-link-type="uri" xlink:href="http://www.sit.puglia.it/portal/portale_rete_ecologica/biomap">http://www.sit.puglia.it/portal/portale_rete_ecologica/biomap</ext-link>), and in yellow, the distribution of <italic>Posidonia oceanica</italic> along the Apulian coastline produced in 2004&#x2013;2005 as part of the project &#x201c;Inventory and Cartography of <italic>Posidonia</italic> Meadows in the Maritime Compartments of Manfredonia, Molfetta, Bari, Brindisi, Gallipoli and Taranto (POR 2000-2006)&#x201d; (<ext-link ext-link-type="uri" xlink:href="https://emodnet.ec.europa.eu/geonetwork/srv/ita/catalog.search#/metadata/e14e1bc8-e52b-4460-b5b3-b5550520728f">https://emodnet.ec.europa.eu/geonetwork/srv/ita/catalog.search#/metadata/e14e1bc8-e52b-4460-b5b3-b5550520728f</ext-link>). In purple, the distribution of mesophotic corals along the Apulian coastline published by Corriero et&#xa0;al. in 2019 (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1038/s41598-019-40284-4">https://doi.org/10.1038/s41598-019-40284-4</ext-link>), and in green, the distribution of the deep oyster reef produced in the framework of the LIFE DREAM Project (<xref ref-type="bibr" rid="B29">Grande et&#xa0;al., 2024</xref>; <ext-link ext-link-type="uri" xlink:href="https://doi.org/10.26383/CNR-ISMAR.2024.6">https://doi.org/10.26383/CNR-ISMAR.2024.6</ext-link>). The area in lilac color is covered by the geomorphological map of the South Adriatic continental margin published in Campiani et&#xa0;al., 2024 (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.1080/17445647.2024.2429707">https://doi.org/10.1080/17445647.2024.2429707</ext-link>), and in orange, the benthic habitat map published by <xref ref-type="bibr" rid="B52">Prampolini et&#xa0;al., 2021</xref> (<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3390/rs13152913">https://doi.org/10.3390/rs13152913</ext-link>). The EMODnet Digital Bathymetry (DTM 2024) provides the background (in blue) and the isobaths (blue lines with 10 meters interval).</p>
</caption></supplementary-material>
<supplementary-material xlink:href="Supplementaryfile1.docx" id="SM1" mimetype="application/vnd.openxmlformats-officedocument.wordprocessingml.document"/></sec>
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