<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD JATS (Z39.96) Journal Publishing DTD v1.3 20210610//EN" "JATS-journalpublishing1-3-mathml3.dtd">
<article xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink" xmlns:ali="http://www.niso.org/schemas/ali/1.0/" xmlns:xsi="http://www.w3.org/2001/XMLSchema-instance" article-type="research-article" dtd-version="1.3" xml:lang="EN">
<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>
</journal-meta>
<article-meta>
<article-id pub-id-type="doi">10.3389/fmars.2026.1769853</article-id>
<article-version article-version-type="Version of Record" vocab="NISO-RP-8-2008"/>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Original Research</subject>
</subj-group>
</article-categories>
<title-group>
<article-title>Integrating benthic ecological status and spatial planning to guide sustainable bottom-contacting fisheries in the Eastern Mediterranean</article-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name><surname>Tsikopoulou</surname><given-names>Irini</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<xref ref-type="corresp" rid="c001"><sup>*</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/625691/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="visualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/visualization/">Visualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Formal analysis" vocab-term-identifier="https://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Project-administration" vocab-term-identifier="https://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
<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>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="conceptualization" vocab-term-identifier="https://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Maina</surname><given-names>Irida</given-names></name>
<xref ref-type="aff" rid="aff2"><sup>2</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/746126/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Papadopoulou</surname><given-names>Nadia</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/170309/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Bastardie</surname><given-names>Fran&#xe7;ois</given-names></name>
<xref ref-type="aff" rid="aff3"><sup>3</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/137808/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="methodology" vocab-term-identifier="https://credit.niso.org/contributor-roles/methodology/">Methodology</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
</contrib>
<contrib contrib-type="author">
<name><surname>Smith</surname><given-names>Chris J.</given-names></name>
<xref ref-type="aff" rid="aff1"><sup>1</sup></xref>
<uri xlink:href="https://loop.frontiersin.org/people/334304/overview"/>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="supervision" vocab-term-identifier="https://credit.niso.org/contributor-roles/supervision/">Supervision</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Data curation" vocab-term-identifier="https://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
<role vocab="credit" vocab-identifier="https://credit.niso.org/" vocab-term="Writing &#x2013; review &amp; editing" vocab-term-identifier="https://credit.niso.org/contributor-roles/writing-review-editing/">Writing &#x2013; review &amp; editing</role>
</contrib>
</contrib-group>
<aff id="aff1"><label>1</label><institution>Institute of Marine Biological Resources and Inland Waters</institution>,&#xa0;<city>Heraklion</city>, <country country="gr">Greece</country></aff>
<aff id="aff2"><label>2</label><institution>Institute of Marine Biological Resources and Inland Waters</institution>,&#xa0;<city>Anavyssos</city>, <country country="gr">Greece</country></aff>
<aff id="aff3"><label>3</label><institution>Section for Ecosystem-Based Management, Technical University of Denmark, National Institute of Aquatic Resources</institution>,&#xa0;<city>Kgs. Lyngby</city>, <country country="dk">Denmark</country></aff>
<author-notes>
<corresp id="c001"><label>*</label>Correspondence: Irini Tsikopoulou, <email xlink:href="mailto:itsikopoulou@hcmr.gr">itsikopoulou@hcmr.gr</email></corresp>
</author-notes>
<pub-date publication-format="electronic" date-type="pub" iso-8601-date="2026-02-27">
<day>27</day>
<month>02</month>
<year>2026</year>
</pub-date>
<pub-date publication-format="electronic" date-type="collection">
<year>2026</year>
</pub-date>
<volume>13</volume>
<elocation-id>1769853</elocation-id>
<history>
<date date-type="received">
<day>17</day>
<month>12</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>13</day>
<month>02</month>
<year>2026</year>
</date>
<date date-type="rev-recd">
<day>09</day>
<month>02</month>
<year>2026</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2026 Tsikopoulou, Maina, Papadopoulou, Bastardie and Smith.</copyright-statement>
<copyright-year>2026</copyright-year>
<copyright-holder>Tsikopoulou, Maina, Papadopoulou, Bastardie and Smith</copyright-holder>
<license>
<ali:license_ref start_date="2026-02-27">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>To inform ecosystem-based fisheries management in line with the EU legislation objectives for marine fisheries, we evaluated the ecological outcomes of alternative spatial management scenarios to fisheries that consider the ecological impact of bottom trawling on the benthic ecosystem in the Eastern Ionian Sea. Trawling intensity in terms of swept area ratio (SAR) and benthic community sensitivity were combined to estimate the relative benthic status (RBS). Then, five management scenarios were tested. The scenarios include four static closure scenarios (below 800 m, below 600 m, the least-trawled 10% of fishing grounds, and areas shallower than 150 m), where trawling is completely excluded without fishing effort redistribution, and one with a trawl ban in all marine protected areas, where fishing effort displacement is modelled dynamically. Baseline RBS was high (&gt;0.9 on a scale of 0 to 1 where 1 is unaffected benthic community) in all habitats, reflecting relatively low benthic degradation due to bottom trawling. Excluding bottom trawling from areas shallower than 150 m in depth produced the greatest improvements, while thresholds at 600 or 800 m depth, and the closure of the 10% least-trawled grounds, had no significant effects on benthic ecological status. Closure of trawling in the marine protected areas produced mixed outcomes, with improvements in some habitats but localized declines due to displaced effort in others. Our study demonstrates the value of including benthic indicators in spatial management strategies to guide adaptive, evidence-based fisheries governance, balancing conservation objectives with socio-economic sustainability.</p>
</abstract>
<kwd-group>
<kwd>benthic ecological status</kwd>
<kwd>bottom trawling impacts</kwd>
<kwd>effort displacement</kwd>
<kwd>EU nature restoration regulation</kwd>
<kwd>fishing closures scenarios</kwd>
<kwd>seafloor integrity</kwd>
<kwd>sustainable management</kwd>
</kwd-group> 
<funding-group>
<award-group id="gs1">
<funding-source id="sp1">
<institution-wrap>
<institution>Horizon 2020 Framework Programme</institution>
<institution-id institution-id-type="doi" vocab="open-funder-registry" vocab-identifier="10.13039/open_funder_registry">10.13039/100010661</institution-id>
</institution-wrap>
</funding-source>
<award-id rid="sp1">101059877, 101000318</award-id>
</award-group>
<funding-statement>The author(s) declared that financial support was received for this work and/or its publication. IT was supported by the GES4SEAS project (Horizon Europe 101059877).</funding-statement>
</funding-group>
<counts>
<fig-count count="5"/>
<table-count count="2"/>
<equation-count count="0"/>
<ref-count count="40"/>
<page-count count="11"/>
<word-count count="6016"/>
</counts>
<custom-meta-group>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Marine Conservation and Sustainability</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 Mediterranean Sea is one of the most heavily exploited marine areas in the world. Due to its semi-enclosed nature and the long history of human use, ecological pressures and socio-economic demands are tightly interlinked (<xref ref-type="bibr" rid="B32">Rick et&#xa0;al., 2020</xref>). Fishing is an important activity throughout the sea, supporting local economies and culture, yet it causes significant pressures on marine ecosystems (<xref ref-type="bibr" rid="B19">Fiorentino and Vitale, 2021</xref>; <xref ref-type="bibr" rid="B18">FAO, 2023</xref>). In the Mediterranean, the activity takes place throughout the region from depths ranging 50&#x2013;1000 m and in 2016 to 2018, it was responsible for 27.0% of Mediterranean landings through just 7.9% of the vessel segment (<xref ref-type="bibr" rid="B16">FAO, 2020</xref>). The 1000 m depth trawling limit was set by the General Fisheries Commission for the Mediterranean and Black Sea (GFCM) and intended to protect deep-sea ecosystems, which are particularly vulnerable to fishing pressure and slow to recover (<xref ref-type="bibr" rid="B11">EC, 2006</xref>). Among fishing methods, bottom trawling is the most pervasive and destructive activity for the seabed due to its direct contact with the benthic environment (<xref ref-type="bibr" rid="B25">Kaiser, 1998</xref>; <xref ref-type="bibr" rid="B20">Foden et&#xa0;al., 2011</xref>). As well as direct removal of benthic and demersal species, bottom trawl gear reshapes benthic community structure, reduces habitat integrity, and affects the biodiversity and overall resilience of marine ecosystems (<xref ref-type="bibr" rid="B13">Eigaard et&#xa0;al., 2016</xref>; <xref ref-type="bibr" rid="B36">Tiano et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B10">Bradshaw et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B1">Agnetta et&#xa0;al., 2025</xref>). In Europe it is the most widespread marine activity with varying levels of intensity and impact across almost all shelf areas (<xref ref-type="bibr" rid="B24">Hiddink et&#xa0;al., 2026</xref>).</p>
<p>Until recently, scientific and policy debates about Mediterranean fisheries focused on whether fish stocks were overexploited and if so how to best reduce this pressure, neglecting the ecological impacts of fishing on benthic habitats (<xref ref-type="bibr" rid="B3">Bahamon et&#xa0;al., 2024</xref>; <xref ref-type="bibr" rid="B5">Bastardie et&#xa0;al., 2025</xref>). For example, the GFCM 2030 Strategy proposes to establish effective area-based conservation measures based on the mapping of essential fish habitats and vulnerable marine ecosystems (VMEs) (<xref ref-type="bibr" rid="B17">FAO, 2021</xref>). However, it pays little or no attention to other benthic habitats that cover most of the seafloor and offer invaluable services to humans, such as carbon sequestration (<xref ref-type="bibr" rid="B29">Porz et&#xa0;al., 2022</xref>). Yet, growing evidence highlights that all benthic ecosystems are highly sensitive to bottom-contacting fishing (<xref ref-type="bibr" rid="B36">Tiano et&#xa0;al., 2019</xref>; <xref ref-type="bibr" rid="B28">Pitcher et&#xa0;al., 2022</xref>). In addition, recent policy and legislation in the European Union has incorporated not only biodiversity but also seafloor integrity as key descriptors of good environmental status (GES in EU, 2008 &#x2013; Marine Strategy Framework Directive, MSFD), placing benthic habitats at the heart of marine sustainability objectives. More recently and in accordance with the MSFD, the Nature Restoration regulation (<xref ref-type="bibr" rid="B14">EU, 2024</xref>) sets specific restoration targets for degraded benthic habitats. In this context, policy around fisheries should also consider seafloor ecological status. To achieve this, mapping the spatial footprint, i.e. the extent and intensity of seafloor disturbance caused by bottom trawls and also assessing benthic status are essential steps for identifying sensitive areas and areas of high pressure, guiding more effective, spatially explicit management strategies (<xref ref-type="bibr" rid="B9">Bona et&#xa0;al., 2025</xref>).</p>
<p>The Eastern Ionian Sea offers a valuable case study for examining the dynamics and trade-offs between bottom trawling and benthic ecological impacts. This region hosts a mosaic of habitats&#x2014;ranging across multiple MSFD broad habitat types from infralittoral to deep bathyal zones, each supporting distinct benthic assemblages with varying sensitivity to disturbance (<xref ref-type="bibr" rid="B35">Smith et&#xa0;al., 2023</xref>). This area includes Patraikos Gulf, one of Greece&#x2019;s main fishing grounds. The Gulf is also adjacent to a marine protected area (MPA), Zakynthos national park, and therefore this area illustrates the tension between ecological protection and extractive fishing industries. Few studies have tested the impact of different spatial management scenarios on reducing the environmental footprint of bottom trawling, mainly focusing on the fisheries themselves, i.e. bycatch, fish stock assessment, nursery-ground protection, and on the economic outcomes (<xref ref-type="bibr" rid="B4">Bastardie et&#xa0;al., 2017</xref>, <xref ref-type="bibr" rid="B5">Bastardie et&#xa0;al., 2025</xref>; <xref ref-type="bibr" rid="B34">Sbrana et&#xa0;al., 2025</xref>), with even less providing evidence on how benthic status is influenced (<xref ref-type="bibr" rid="B6">Bastardie et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B40">Zupa et&#xa0;al., 2025</xref>).</p>
<p>In this context, our study aims to evaluate the ecological outcomes of various management measures and fishing restrictions on the benthic ecosystem and to explore the implications of these findings for regional fisheries governance. This is in accordance with the objectives of the EU Biodiversity Strategy to achieve GES in at least 30% of the habitats in the EU marine waters by 2030 (<xref ref-type="bibr" rid="B12">EC, 2021</xref>) and aligns with the recent goals of the GFCM, which among others promotes sustainable fisheries through the protection of vulnerable marine ecosystems (<xref ref-type="bibr" rid="B17">FAO, 2021</xref>). By integrating ecological assessments with management scenarios, this research contributes to the advancement of ecosystem-based fisheries management in the Mediterranean. It provides insights into the trade-offs and co-benefits between marine habitat conservation and resource exploitation, offering evidence-based guidance for environmental and fisheries policymakers seeking to align marine ecosystem health with socio-economic sustainability. This approach has been to anticipate the impacts of different protective management strategies by modelling scenarios of alternative spatial allocations of bottom trawling effort on seabed ecological status.</p>
</sec>
<sec id="s2" sec-type="materials|methods">
<label>2</label>
<title>Materials and methods</title>
<sec id="s2_1">
<label>2.1</label>
<title>Study area</title>
<p>The study area is in the Eastern Ionian Sea in the GFCM geographical sub-area 20 (GSA20) (<xref ref-type="fig" rid="f1"><bold>Figure&#xa0;1</bold></xref>). The area was divided into grid cells of 0.05x0.05 decimal degrees (dd) using the C-square grid approach (<xref ref-type="bibr" rid="B31">Rees, 2003</xref>). The EMODnet seabed habitat data portal (<xref ref-type="bibr" rid="B38">Vasquez et al., 2023</xref>) was used to extract the distribution of MSFD broad habitat types within the study area. In order to link habitat information to fishing intensity, habitat types were transferred into the midpoints of the C-square grid cells. The MSFD broad habitat type that overlapped the central point of the C-square was assigned to the whole C-square. Due to the centroid based assignment as well as the grid resolution, minor spatial resolution artifacts may occur where a single cell spans across bathymetric contours.</p>
<fig id="f1" position="float">
<label>Figure&#xa0;1</label>
<caption>
<p>Bathymetry (left) and map of the MSFD broad habitat types (right) present in the GSA20 [data sourced from <xref ref-type="bibr" rid="B38">Vasquez et al. (2023)</xref>]. Na: no data on habitat type.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-13-1769853-g001.tif">
<alt-text content-type="machine-generated">Side-by-side maps of the western Greek coastline depict marine data for GSA 20. The left map shows bathymetric depth zones ranging from zero to above one thousand meters in varying blue shades. The right map displays MSFD habitat types in multiple colors, such as infralittoral sand, circalittoral mud, and abyssal, with a legend indicating each type. An inset in the lower right provides regional context of GSA 20 in the Mediterranean.</alt-text>
</graphic></fig>
</sec>
<sec id="s2_2">
<label>2.2</label>
<title>Baseline benthic ecological status assessment</title>
<p>To assess the baseline conditions of the benthic ecosystem in the study area, we used the Relative Benthic Status (RBS) indicator proposed by <xref ref-type="bibr" rid="B27">Pitcher et&#xa0;al. (2017)</xref>. The calculation of RBS requires a benthic community sensitivity spatial layer, distribution of fishing intensity layer per type of fishing activity (here bottom trawling), depletion rates of benthic animals affected by specific fishing activity (i.e. m&#xe9;tier) and recovery rates of the benthic community associated with the habitat types (see details in <xref ref-type="bibr" rid="B23">Hiddink et&#xa0;al., 2017</xref>; <xref ref-type="bibr" rid="B27">Pitcher et&#xa0;al., 2017</xref>). RBS takes values from zero to one with higher values indicating better benthic status.</p>
<sec id="s2_2_1">
<label>2.2.1</label>
<title>Benthic community sensitivity assessment</title>
<p>RBS assumes that the sensitivity of the benthic community to bottom trawling is related to the median expected longevity of the community. A community longevity is defined as the combination of the theoretical life expectancy of all the species that make up this community (<xref ref-type="bibr" rid="B33">Rijnsdorp et&#xa0;al., 2020</xref>). The median longevity of the community is the longevity defined at the 50<sup>th</sup> percentile of the cumulative biomass of this community. Benthic community sensitivity was quantified using species longevity traits, a widely applied ecological proxy for vulnerability to bottom trawling disturbance. Briefly, the method required firstly that the macrofauna species from undisturbed stations are assigned to one of four longevity classes (&lt;1 year, 1&#x2013;3 years, 3&#x2013;10 years, &gt;10 years) using a fuzzy-coding approach. Then, the relationship between cumulative biomass and longevity was estimated using generalized linear mixed models (GLMMs) with a binomial distribution. Environmental predictors included bathymetry (log-transformed) and MSFD broad habitat type as fixed effects, while sampling site was included as a random effect. All model combinations were compared using the Akaike Information Criterion (AIC). Benthic sensitivity of the study area was extracted from <xref ref-type="bibr" rid="B35">Smith et&#xa0;al. (2023)</xref>. Sensitivity was available only for specific MSFD broad habitat types, including infralittoral sand, infralittoral mud, circalittoral sand, circalittoral mud, offshore circalittoral mud and upper bathyal sediment. Median longevity was extracted as the indicator of benthic sensitivity for each habitat type in the GSA20.</p>
</sec>
<sec id="s2_2_2">
<label>2.2.2</label>
<title>Depletion and recovery rates</title>
<p>Demersal otter trawls are the only m&#xe9;tier that operates in the study area. The depletion rate for otter trawls was obtained from the global meta-analysis conducted by <xref ref-type="bibr" rid="B23">Hiddink et&#xa0;al. (2017)</xref> and was set at 0.06. Similarly, the recovery rate (<italic>r</italic>) was based on a meta-analysis, calculated as <italic>r = 5.31/longevity</italic> (<xref ref-type="bibr" rid="B22">Hiddink et&#xa0;al., 2019</xref>).</p>
</sec>
<sec id="s2_2_3">
<label>2.2.3</label>
<title>Fishing intensity assessment</title>
<p>Fishing intensity was quantified using the swept area ratio (SAR), an indicator used widely and defined as the area swept by trawl gear relative to the surface area of a grid cell (<xref ref-type="bibr" rid="B2">Amoroso et&#xa0;al., 2018</xref>). Vessel Monitoring System (VMS) data from 2018&#x2013;2020 were used to estimate SAR for demersal otter trawls. Annual SAR values were averaged over 2018&#x2013;2020 to produce a baseline measure of trawling intensity in GSA20. Grid cells were categorized by trawling frequency, and their overlap with habitat types was mapped. Raw VMS data for the years 2018&#x2013;2020 were provided by the Hellenic Ministry of Mercantile Marine and Island Policy and were analyzed based on the methods and specifications further described in <xref ref-type="bibr" rid="B26">Maina et&#xa0;al. (2021)</xref>.</p>
</sec>
</sec>
<sec id="s2_3">
<label>2.3</label>
<title>Fisheries management scenarios</title>
<p>The RBS was calculated for GSA20 by combining the trawling intensity (SAR) with benthic sensitivity (<xref ref-type="bibr" rid="B27">Pitcher et&#xa0;al., 2017</xref>). Since the benthic sensitivity layer was available only for specific habitat types (<xref ref-type="bibr" rid="B35">Smith et&#xa0;al., 2023</xref>), RBS was not calculated for infralittoral biogenic habitats, bathyal and abyssal zone. Five spatial management scenarios were tested to predict their effect on benthic ecological status (i.e. RBS). Scenarios were run by changing SAR according to different rules and recalculating RBS over the area. Scenarios were chosen from a range of possibilities from strict conservation actions to more gentle actions that reflect closing areas of minimal fishing use.</p>
<p>Four of the management scenarios that were tested were based on a reduction of fishing (static approach), i.e. with fishing effort totally cut from closed areas, assuming effort is removed completely. The fifth scenario was dynamic including re-allocation of fishing effort (see section: Specifications for SAR inputs per scenario). The dynamic scenario is expected to provide a more realistic representation of fleet responses, since fishing effort is unlikely to be removed entirely unless authorities specifically reduce effort through spatial management measures. Moreover, given that the focus of this work was to investigate potential changes in RBS, the effects of such management options on alternative resources and on the fisheries economy require further investigation. The following spatial management scenarios were tested:</p>
<list list-type="order">
<list-item>
<p>Scenario 1 (static): Restricting bottom trawling below 800 m, reducing depth of the current Mediterranean-wide 1000 m depth limit of trawling, with no effort displacement. This scenario was inspired by GFCM recommendations for the protection of VMEs.</p></list-item>
<list-item>
<p>Scenario 2 (static): Restricting bottom trawling below 600 m depth, with no effort displacement. This is a stricter version of scenario 1 that was inspired by GFCM recommendations for the protection of VMEs and sensitive habitats.</p></list-item>
<list-item>
<p>Scenario 3 (static): Closure of the least-trawled 10% of core fishing grounds (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2B</bold></xref>), retaining the core fishing areas to attain the least socio-economic consequences. This scenario was inspired by the International Council for the Exploration of the Sea (ICES) as the one with the smallest economic impacts on fisheries.</p></list-item>
<list-item>
<p>Scenario 4 (static): Restricting bottom trawling down to 150 m depth, based on an Oceana proposal<xref ref-type="fn" rid="fn1"><sup>1</sup></xref>. This scenario, although might be perceived as unrealistic due to its potential economic consequences, will effectively exclude from bottom trawling, shallow sensitive habitats such as seagrass meadows, coralligenous reefs, and maerl beds by creating broad-scale trawl-free areas in the coastal zone.</p></list-item>
<list-item>
<p>Scenario 5 (dynamic): Restricting bottom trawling in all Eastern Ionian Sea designated MPAs (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2A</bold></xref>), including National Parks and Reserves, RAMSAR Sites and other Special Sites/Areas and Refuges, with effort redistributed. This scenario was inspired by the EU Action Plan for protecting and restoring marine ecosystems to achieve sustainable and resilient fisheries (<xref ref-type="bibr" rid="B15">European Commission, 2023</xref>). EU Member States are called on, among other things &#x201c;to adopt national measures and, where relevant, submit joint recommendations to the Commission to ensure that mobile bottom fishing is phased out, in all MPAs by 2030&#x201d;.</p></list-item>
</list>
<fig id="f2" position="float">
<label>Figure&#xa0;2</label>
<caption>
<p><bold>(A)</bold> Distribution of the MPAs and other protected areas in the study area. MPAs within GSA20 include National Parks, National Marine Parks, National Reserve Areas, Ramsar sites, Wildlife Refuges, Sites of Community Importances, Special Areas of Conservation and Special Protection Areas. <bold>(B)</bold> Distribution of the 10% least-trawled fishing grounds in the study area.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-13-1769853-g002.tif">
<alt-text content-type="machine-generated">Panel A shows a coastal map with orange areas representing marine protected areas along the shoreline. Panel B displays the same coast with blue squares indicating the ten percent least-trawled fishing grounds.</alt-text>
</graphic></fig>
</sec>
<sec id="s2_4">
<label>2.4</label>
<title>Specifications for SAR inputs per scenario</title>
<p>For Scenarios 1&#x2013;4 (static scenarios), SAR within closed areas was set to zero (SAR = 0 in all C-squares inside banned zones) assuming trawling effort is removed completely from the area. In all other C-squares, SAR was based on mean values for 2018&#x2013;2020.</p>
<p>For Scenario 5 (dynamic scenario), SAR values were obtained from simulations using the DISPLACE spatially explicit bioeconomic model (<xref ref-type="bibr" rid="B7">Bastardie et&#xa0;al., 2014</xref>; <ext-link ext-link-type="uri" xlink:href="http://www.displace-project.org">www.displace-project.org</ext-link>). The DISPLACE framework simulates the behavior of individual fishing vessels by integrating fleet dynamics, resource availability, and fisheries management options. It enables the evaluation of biological, ecosystem, economic, and energy efficiency effects resulting from the redistribution of fishing effort to alternative grounds. The Eastern Ionian Sea application was parameterized with data on fishing effort distribution, fish stock traits, species abundance, selectivity, and economic indicators (see details in <xref ref-type="bibr" rid="B26">Maina et&#xa0;al., 2021</xref>; <xref ref-type="bibr" rid="B5">Bastardie et&#xa0;al., 2025</xref>). Fishing effort redistribution was assumed to occur within GSA 20, reflecting the typical behavior of the local fleet. The model can also incorporate benthic community dynamics by coupling gear-specific depletion rates with habitat-specific, trait-based recovery rates (<xref ref-type="bibr" rid="B6">Bastardie et&#xa0;al., 2020</xref>). However, this benthic component is still under development for the Eastern Ionian Sea application. Simulations performed for the period 2020&#x2013;2027 aimed at evaluating the effects of effort displacement from all MPAs, to alternative fishing grounds. The simulated SAR was derived from 50 Monte Carlo replicates, allowing estimation of uncertainty around the mean SAR. The simulated SAR values (at C-square resolution) were used for Scenario 1 purposes and correspond to the final simulated year (2027). The resulting SAR was then combined with benthic sensitivity data to estimate Relative Benthic Status (RBS).</p>
</sec>
<sec id="s2_5">
<label>2.5</label>
<title>Estimation of RBS</title>
<p>RBS was calculated for all management scenarios and compared with the baseline (status quo) conditions, i.e., the mean RBS for the period 2018&#x2013;2022. A threshold of RBS &gt; 0.8 was used as a candidate benchmark for Good Environmental Status (<xref ref-type="bibr" rid="B35">Smith et&#xa0;al., 2023</xref>). The non-parametric Wilcoxon signed-rank test was selected to compare RBS between baseline conditions and the different management scenarios per habitat type (<xref ref-type="bibr" rid="B21">Haynes, 2013</xref>), since the normality and homogeneity of variance assumptions were violated. Statistical significance was determined at a threshold of p-value &lt; 0.05.</p>
<p>RBS estimations were carried out in the R software (<xref ref-type="bibr" rid="B30">R Core Team, 2024</xref>). Scripts were developed by adapting the code from <xref ref-type="bibr" rid="B37">van Denderen et&#xa0;al. (2020)</xref>.</p>
</sec>
</sec>
<sec id="s3" sec-type="results">
<label>3</label>
<title>Results</title>
<sec id="s3_1">
<label>3.1</label>
<title>Baseline conditions</title>
<p>Baseline fishing pressure exceeded SAR values of 1 only in 17% of the trawled cells in the grid (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3a</bold></xref>). Maximum SAR value recorded in the area was approximately 2.8. This value was recorded in only one grid cell emphasizing that trawling pressure did not exceed SAR = 3 even in areas of concentrated trawling. The baseline RBS reflected the annual average trawling pressure distribution in GSA20 from 2018-2020. RBS was higher than the 0.8 threshold, that defines GES, in the entire study area, indicating minimal impact from bottom trawling. The lowest RBS values (still &gt;0.8) were recorded in the centre of the study area (outer Patraikos Gulf) where the main fishing grounds are found &#x2013; with highest SAR intensities (<xref ref-type="fig" rid="f3"><bold>Figure&#xa0;3b</bold></xref>). When aggregated by MSFD broad habitat type, RBS values exceeded 0.9 in all cases indicating low average impacts across the assessed habitats (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). The most impacted habitat for the period 2018&#x2013;2020 was offshore circalittoral mud where more than 85% of the area that this habitat occupies was bottom trawled. Circalittoral sand exhibited the highest average SAR values (0.76), followed by offshore circalittoral mud (0.60) (<xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). In contrast, infralittoral and bathyal sediments showed SAR values close to zero due to European and national legislation that prohibit trawling in shallow (i.e. within 3 nm distance from the coast or less than 50 m depth) and deeper depths (i.e. below 1000 m depth) as well as in certain areas/gulfs (see <xref ref-type="bibr" rid="B26">Maina et&#xa0;al., 2021</xref>).</p>
<fig id="f3" position="float">
<label>Figure&#xa0;3</label>
<caption>
<p><bold>(A)</bold> Swept area ratio (SAR) and <bold>(B)</bold> baseline benthic conditions as reflected by the RBS indicator in the study area for the period 2018-2020.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-13-1769853-g003.tif">
<alt-text content-type="machine-generated">Side-by-side maps showing the same coastal region with hexagonal grids. Panel A uses a red gradient to indicate Swept Area Ratio (SAR) from less than 0.5 to 2.5–3. Panel B uses yellow-to-red shades for Relative Benthic Status (RBS) from 0.7–0.8 up to 0.95–1, and pale yellow for areas with no trawling. Land areas are shown in gray.</alt-text>
</graphic></fig>
<table-wrap id="T1" position="float">
<label>Table&#xa0;1</label>
<caption>
<p>Area, SAR, proportion of trawled cells and RBS estimated per MSFD broad habitat type in the GSA20 estimated for the period 2018-2020 (baseline) and under different spatial management scenarios.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" rowspan="3" align="center">MSFD broad habitat types</th>
<th valign="middle" rowspan="3" align="center">area (x 10<sup>3</sup> km<sup>2</sup>)</th>
<th valign="middle" rowspan="2" colspan="3" align="center"><italic>Baseline conditions</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Scenario 1</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Scenario 2</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Scenario 3</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Scenario 4</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Scenario 5</italic></th>
</tr>
<tr>
<th valign="middle" colspan="3" align="center"><italic>Restricting bottom trawling below 800 m</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Restricting bottom trawling below 600 m</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Closure of the least-trawled 10% of core fishing grounds</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Restricting bottom trawling down to 150 m</italic></th>
<th valign="middle" colspan="3" align="center"><italic>Restricting bottom trawling in all MPAs</italic></th>
</tr>
<tr>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
<th valign="middle" align="center">SAR</th>
<th valign="middle" align="center">% of cells trawled</th>
<th valign="middle" align="center">RBS</th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="right">Infralittoral sand</td>
<td valign="middle" align="right">1.75</td>
<td valign="middle" align="right">0.038</td>
<td valign="middle" align="right">9.59</td>
<td valign="middle" align="right">0.968</td>
<td valign="middle" align="right">0.038</td>
<td valign="middle" align="right">9.59</td>
<td valign="middle" align="right">0.9679</td>
<td valign="middle" align="right">0.0378</td>
<td valign="middle" align="right">9.59</td>
<td valign="middle" align="right">0.9679</td>
<td valign="middle" align="right">0.037</td>
<td valign="middle" align="right">8.22</td>
<td valign="middle" align="right">0.9679</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0.035</td>
<td valign="middle" align="right">6.85</td>
<td valign="middle" align="right">0.9681</td>
</tr>
<tr>
<td valign="middle" align="right">Infralittoral mud</td>
<td valign="middle" align="right">0.27</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">0.001</td>
<td valign="middle" align="right">9.09</td>
<td valign="middle" align="right">0.9999</td>
</tr>
<tr>
<td valign="middle" align="right">Infralittoral rock and biogenic reef</td>
<td valign="middle" align="right">0.10</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">-</td>
</tr>
<tr>
<td valign="middle" align="right">Circalittoral sand</td>
<td valign="middle" align="right">2.02</td>
<td valign="middle" align="right">0.759</td>
<td valign="middle" align="right">64.29</td>
<td valign="middle" align="right">0.971</td>
<td valign="middle" align="right">0.757</td>
<td valign="middle" align="right">63.10</td>
<td valign="middle" align="right">0.9714</td>
<td valign="middle" align="right">0.757</td>
<td valign="middle" align="right">63.10</td>
<td valign="middle" align="right">0.9714</td>
<td valign="middle" align="right">0.758</td>
<td valign="middle" align="right">60.71</td>
<td valign="middle" align="right">0.9714</td>
<td valign="middle" align="right">0.007</td>
<td valign="middle" align="right">4.76</td>
<td valign="middle" align="right">0.9997</td>
<td valign="middle" align="right">0.654</td>
<td valign="middle" align="right">58.33</td>
<td valign="middle" align="right">0.9752</td>
</tr>
<tr>
<td valign="middle" align="right">Circalittoral mud</td>
<td valign="middle" align="right">0.63</td>
<td valign="middle" align="right">0.552</td>
<td valign="middle" align="right">55.56</td>
<td valign="middle" align="right">0.986</td>
<td valign="middle" align="right">0.552</td>
<td valign="middle" align="right">55.56</td>
<td valign="middle" align="right">0.9856</td>
<td valign="middle" align="right">0.552</td>
<td valign="middle" align="right">55.56</td>
<td valign="middle" align="right">0.9856</td>
<td valign="middle" align="right">0.552</td>
<td valign="middle" align="right">51.85</td>
<td valign="middle" align="right">0.9857</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">1</td>
<td valign="middle" align="right">1.073</td>
<td valign="middle" align="right">51.85</td>
<td valign="middle" align="right">0.9726</td>
</tr>
<tr>
<td valign="middle" align="right">Offshore circalittoral sand</td>
<td valign="middle" align="right">0.68</td>
<td valign="middle" align="right">0.249</td>
<td valign="middle" align="right">57.14</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0.249</td>
<td valign="middle" align="right">57.14</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0.249</td>
<td valign="middle" align="right">57.14</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0.246</td>
<td valign="middle" align="right">50.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0.039</td>
<td valign="middle" align="right">25.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0.103</td>
<td valign="middle" align="right">42.86</td>
<td valign="middle" align="right">&#x2013;</td>
</tr>
<tr>
<td valign="middle" align="right">Offshore circalittoral mud</td>
<td valign="middle" align="right">0.58</td>
<td valign="middle" align="right">0.608</td>
<td valign="middle" align="right">87.50</td>
<td valign="middle" align="right">0.980</td>
<td valign="middle" align="right">0.608</td>
<td valign="middle" align="right">87.50</td>
<td valign="middle" align="right">0.9798</td>
<td valign="middle" align="right">0.608</td>
<td valign="middle" align="right">87.50</td>
<td valign="middle" align="right">0.9798</td>
<td valign="middle" align="right">0.606</td>
<td valign="middle" align="right">83.33</td>
<td valign="middle" align="right">0.9798</td>
<td valign="middle" align="right">0.135</td>
<td valign="middle" align="right">45.83</td>
<td valign="middle" align="right">0.9956</td>
<td valign="middle" align="right">1.027</td>
<td valign="middle" align="right">79.17</td>
<td valign="middle" align="right">0.9656</td>
</tr>
<tr>
<td valign="middle" align="right">Upper bathyal sediment</td>
<td valign="middle" align="right">4.11</td>
<td valign="middle" align="right">0.110</td>
<td valign="middle" align="right">62.72</td>
<td valign="middle" align="right">0.997</td>
<td valign="middle" align="right">0.110</td>
<td valign="middle" align="right">62.72</td>
<td valign="middle" align="right">0.9965</td>
<td valign="middle" align="right">0.107</td>
<td valign="middle" align="right">60.95</td>
<td valign="middle" align="right">0.9965</td>
<td valign="middle" align="right">0.109</td>
<td valign="middle" align="right">49.11</td>
<td valign="middle" align="right">0.9966</td>
<td valign="middle" align="right">0.073</td>
<td valign="middle" align="right">53.25</td>
<td valign="middle" align="right">0.9976</td>
<td valign="middle" align="right">0.081</td>
<td valign="middle" align="right">41.42</td>
<td valign="middle" align="right">0.9974</td>
</tr>
<tr>
<td valign="middle" align="right">Lower bathyal sediment</td>
<td valign="middle" align="right">104.48</td>
<td valign="middle" align="right">0.003</td>
<td valign="middle" align="right">1.82</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.003</td>
<td valign="middle" align="right">1.31</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.001</td>
<td valign="middle" align="right">0.77</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.003</td>
<td valign="middle" align="right">0.84</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.003</td>
<td valign="middle" align="right">1.82</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.002</td>
<td valign="middle" align="right">0.58</td>
<td valign="middle" align="right">-</td>
</tr>
<tr>
<td valign="middle" align="right">Abyssal</td>
<td valign="middle" align="right">3.26</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
<td valign="middle" align="right">0</td>
<td valign="middle" align="right">0.00</td>
<td valign="middle" align="right">&#x2013;</td>
</tr>
<tr>
<td valign="middle" align="right">Unknown</td>
<td valign="middle" align="right">4.40</td>
<td valign="middle" align="right">0.154</td>
<td valign="middle" align="right">32.29</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.151</td>
<td valign="middle" align="right">30.21</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.147</td>
<td valign="middle" align="right">28.65</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.153</td>
<td valign="middle" align="right">29.69</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.021</td>
<td valign="middle" align="right">10.42</td>
<td valign="middle" align="right">-</td>
<td valign="middle" align="right">0.161</td>
<td valign="middle" align="right">18.75</td>
<td valign="middle" align="right">-</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s3_2">
<label>3.2</label>
<title>Scenario 1: restricting bottom trawling below 800 m depth (static)</title>
<p>This scenario excluded 7% of the baseline trawled area and produced no significant changes from the baseline conditions (<xref ref-type="table" rid="T1"><bold>Tables&#xa0;1</bold></xref>, <xref ref-type="table" rid="T2"><bold>2</bold></xref>). RBS values across most habitats remained identical to baseline estimates, as trawling activity below 1000 m depth was already prohibited (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4a</bold></xref>). The proportion of trawled cells decreased slightly for lower bathyal sediments, with no measurable change at 4 decimal precision points in the RBS.</p>
<table-wrap id="T2" position="float">
<label>Table&#xa0;2</label>
<caption>
<p>Wilcoxon signed-rank test results comparing RBS indicators between baseline conditions and each scenario.</p>
</caption>
<table frame="hsides">
<thead>
<tr>
<th valign="middle" align="center"/>
<th valign="middle" align="center"/>
<th valign="middle" colspan="4" align="center"><italic>Scenario 1</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Scenario 2</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Scenario 3</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Scenario 4</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Scenario 5</italic></th>
</tr>
<tr>
<th valign="middle" align="center"/>
<th valign="middle" align="center"/>
<th valign="middle" colspan="4" align="center"><italic>Restricting bottom trawling below 800 m</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Restricting bottom trawling below 600 m</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Closure of the least-trawled 10% of core fishing grounds</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Restricting bottom trawling down to 150 m</italic></th>
<th valign="middle" colspan="4" align="center"><italic>Restricting bottom trawling in all MPAs</italic></th>
</tr>
<tr>
<th valign="middle" align="center"><italic>MSFD broad habitat types</italic></th>
<th valign="middle" align="center"><italic>Sample size</italic></th>
<th valign="middle" align="center"><italic>V-statistic</italic></th>
<th valign="middle" align="center"><italic>p-value</italic></th>
<th valign="middle" align="center"><italic>Significance</italic></th>
<th valign="middle" align="center"><italic>Change in RBS</italic></th>
<th valign="middle" align="center"><italic>V-statistic</italic></th>
<th valign="middle" align="center"><italic>p-value</italic></th>
<th valign="middle" align="center"><italic>Significance</italic></th>
<th valign="middle" align="center"><italic>Change in RBS</italic></th>
<th valign="middle" align="center"><italic>V-statistic</italic></th>
<th valign="middle" align="center"><italic>p-value</italic></th>
<th valign="middle" align="center"><italic>Significance</italic></th>
<th valign="middle" align="center"><italic>Change in RBS</italic></th>
<th valign="middle" align="center"><italic>V-statistic</italic></th>
<th valign="middle" align="center"><italic>p-value</italic></th>
<th valign="middle" align="center"><italic>Significance</italic></th>
<th valign="middle" align="center"><italic>Change in RBS</italic></th>
<th valign="middle" align="center"><italic>V-statistic</italic></th>
<th valign="middle" align="center"><italic>p-value</italic></th>
<th valign="middle" align="center"><italic>Significance</italic></th>
<th valign="middle" align="center"><italic>Change in RBS</italic></th>
</tr>
</thead>
<tbody>
<tr>
<td valign="middle" align="right">Infralittoral sand</td>
<td valign="middle" align="center">73</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0.02</td>
<td valign="middle" align="center">*</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">6</td>
<td valign="middle" align="center">0.2</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
</tr>
<tr>
<td valign="middle" align="right">Infralittoral mud</td>
<td valign="middle" align="center">11</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
</tr>
<tr>
<td valign="middle" align="right">Circalittoral sand</td>
<td valign="middle" align="center">84</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0.18</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">123</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
</tr>
<tr>
<td valign="middle" align="right">Circalittoral mud</td>
<td valign="middle" align="center">27</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">93</td>
<td valign="middle" align="center">0.06</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
</tr>
<tr>
<td valign="middle" align="right">Offshore circalittoral mud</td>
<td valign="middle" align="center">24</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">1</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0.01</td>
<td valign="middle" align="center">**</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">118</td>
<td valign="middle" align="center">0.94</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
</tr>
<tr>
<td valign="middle" align="right">Upper bathyal sediment</td>
<td valign="middle" align="center">169</td>
<td valign="middle" align="center">3</td>
<td valign="middle" align="center">0.37</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">55</td>
<td valign="middle" align="center">0.53</td>
<td valign="middle" align="center">ns</td>
<td valign="middle" align="center"/>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
<td valign="middle" align="center">1610</td>
<td valign="middle" align="center">0</td>
<td valign="middle" align="center">***</td>
<td valign="middle" align="center">Improvement</td>
</tr>
</tbody>
</table>
<table-wrap-foot>
<fn>
<p>ns, not significant; *p &lt; 0.05; **p&lt;0.01, ***p&lt;0.001.</p></fn>
</table-wrap-foot>
</table-wrap>
<fig id="f4" position="float">
<label>Figure&#xa0;4</label>
<caption>
<p>Predicted benthic conditions in the study area as reflected by the RBS indicator under the static scenarios: <bold>(a)</bold> Scenario 1: Restricting bottom trawling below 800 m, <bold>(b)</bold> Scenario 2: Restricting bottom trawling below 600 m, <bold>(c)</bold> Scenario 3: Closure of the least-trawled 10% of core fishing grounds, <bold>(d)</bold> Scenario 4: Restricting bottom trawling down to 150 m.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-13-1769853-g004.tif">
<alt-text content-type="machine-generated">Four-panel geographic heatmap shows the relative benthic status (RBS) of an area in Greece, with color coding from red (0.7-0.8) to yellow (0.95-1) and gray for land, across panels a, b, c, and d. Each panel displays similar spatial patterns of RBS values, with highest disturbance around latitude 38 degrees north and longitude 21 degrees east, and less disturbance further from this area. A legend at bottom right explains the RBS color scale and designates “no trawling” in pale yellow.</alt-text>
</graphic></fig>
</sec>
<sec id="s3_3">
<label>3.3</label>
<title>Scenario 2: restricting bottom trawling below 600 m depth (static)</title>
<p>Similar to the 800 m scenario, the 600 m depth restriction excluded 16% of the trawled area but produced no substantial improvements in RBS values per habitat type (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4b</bold></xref>; <xref ref-type="table" rid="T1"><bold>Tables&#xa0;1</bold></xref>, <xref ref-type="table" rid="T2"><bold>2</bold></xref>). The reduction in the proportion of trawled cells was slightly greater than under Scenario 2, but overall differences compared to the baseline remained minimal. In all habitat types, RBS values were above 0.97, with upper bathyal sediments showing no significant increase in benthic status (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>).</p>
</sec>
<sec id="s3_4">
<label>3.4</label>
<title>Scenario 3: closure of the least-trawled 10% of core fishing grounds (static)</title>
<p>Closing the least-trawled 10% of core fishing grounds although excluded 22% of the baseline trawled area, had a minor effect on the benthic status (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4c</bold></xref>). SAR and RBS remained nearly unchanged across all habitats compared to the baseline conditions although in the upper bathyal sediment the increase in RBS was statistically highly significant (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>).</p>
</sec>
<sec id="s3_5">
<label>3.5</label>
<title>Scenario 4: restricting bottom trawling down to 150 m (static)</title>
<p>The most substantial changes were observed under Scenario 4, which prohibited trawling down to 150 m depth, excluding 41% of the baseline trawled area (<xref ref-type="fig" rid="f4"><bold>Figure&#xa0;4d</bold></xref>; <xref ref-type="table" rid="T1"><bold>Table&#xa0;1</bold></xref>). RBS in all habitat types, except for the infralittoral zone, significantly increased (<xref ref-type="table" rid="T2"><bold>Table&#xa0;2</bold></xref>). SAR values for these habitats dropped to zero, and the proportion of trawled cells was reduced correspondingly.</p>
</sec>
<sec id="s3_6">
<label>3.6</label>
<title>Scenario 5: restricting bottom trawling in all MPAs (dynamic)</title>
<p>Among the spatial management scenarios, dynamic Scenario 5 restricted trawling in all protected areas (<xref ref-type="fig" rid="f2"><bold>Figure&#xa0;2A</bold></xref>), excluding 36% of the baseline trawled area. This resulted in statistically significant increase (improvement) in RBS values for some habitat types, such as in the circalittoral sand and the upper bathyal sediments (<xref ref-type="table" rid="T1"><bold>Tables&#xa0;1</bold></xref>, <xref ref-type="table" rid="T2"><bold>2</bold></xref>). However, redistribution of fishing effort to other nearby grounds led to localized decreases in RBS, most notably in Patraikos Gulf, where fishing effort was concentrated (<xref ref-type="fig" rid="f5"><bold>Figure&#xa0;5</bold></xref>). Although a decrease in average RBS was recorded in the offshore circalittoral mud, it was not statistically significant. It is also worth noting that although the average benthic status in all habitat types was above 0.8, there is a certain area in the inner Patraikos Gulf where the RBS was below this threshold (<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>Predicted benthic conditions as reflected in RBS indicator under the dynamic Scenario 5 in the study area.</p>
</caption>
<graphic mimetype="image" mime-subtype="tiff" xlink:href="fmars-13-1769853-g005.tif">
<alt-text content-type="machine-generated">Heatmap showing trawling activity across latitudes 36°N to 39°N and longitudes 20°E to 23°E, with color-coded Relative Benthic Status (RBS) ranging from 0.7 to 1 and areas of no trawling.</alt-text>
</graphic></fig>
</sec>
</sec>
<sec id="s4" sec-type="discussion">
<label>4</label>
<title>Discussion</title>
<p>Mapping regional trawling activity and benthic ecological status are valuable policy tools, enabling effective spatial management by identifying sensitive or marine areas at high risk to disturbance (<xref ref-type="bibr" rid="B9">Bona et&#xa0;al., 2025</xref>). Insufficient knowledge of fisheries distribution and sensitive areas can lead to incorrectly placed closures that will not only result in a reduction of economic profit but may also fail to provide benefits to maintain or improve the benthic ecological status (<xref ref-type="bibr" rid="B39">Vaughan, 2017</xref>; <xref ref-type="bibr" rid="B34">Sbrana et&#xa0;al., 2025</xref>). Based on the benthic status (RBS) estimated with recent trawling effort in the area by <xref ref-type="bibr" rid="B28">Pitcher et&#xa0;al. (2022)</xref> and <xref ref-type="bibr" rid="B35">Smith et&#xa0;al. (2023)</xref>, this study explores how different fisheries spatial management scenarios would change RBS. Trawling intensity and benthic sensitivity are linked across different habitat types and alternative management measures. Our findings suggest that the selection of specific management measures is critical, since different measures have yielded different outcomes. While bottom trawling closures in shallow waters (&lt;150m) offered the most substantial improvement in RBS, deeper bathymetric closures and the displacement of effort from protected areas resulted in negligible or mixed effects.</p>
<p>Techniques for reconciling fisheries&#x2019; economic performance and minimization of their impacts on the seafloor are of growing interest to policymakers now that environmental targets are pursued (within the GFCM, the MSFD, and the Nature Restoration Regulation). In this context, this study aimed to evaluate the performance of different management scenarios to reduce the ecological impact of bottom trawling on the seafloor. This study prioritized seafloor integrity in the management scenarios tested, with the expectation that preserving benthic habitats will, in turn, promote the long-term sustainability of the fisheries sector. Among the scenarios examined, four of them were based on a bathymetric ban of trawling where fishing pressure is removed from the system entirely. The outcomes of these scenarios indicated that reducing trawling effort does not always improve benthic status. Specifically, restricting trawling down to 150 m depth yielded the strongest improvements, while closures of trawling in areas deeper than 600 or 800 m had negligible changes in the benthic status. The latter has been suggested as a sustainable measure that would allow the protection of the deep-sea including several deep-water coral species and vulnerable marine ecosystems, with relatively little economic impact (<xref ref-type="bibr" rid="B34">Sbrana et&#xa0;al., 2025</xref>). Nevertheless, the negligible difference in RBS between the 800 m and 600 m scenarios suggests that for the Eastern Ionian Sea, the &#x2018;ecological gain&#x2019; is relatively small, indicating that targeted, habitat-specific interventions rather than broad bathymetric shifts may be more beneficial. In contrast, the implementation of a drastic management plan that would offer protection to the shallower habitats (to 150 m depth) delivered significant ecological benefits in the model, first explained as these habitats host the core fishing grounds in the area and also host the most sensitive benthic communities (<xref ref-type="bibr" rid="B35">Smith et&#xa0;al., 2023</xref>). However, as maximum trawling intensity is generally observed at 50&#x2013;200 m depth, this scenario may have significant economic costs to fishers, particularly if no &#x201c;reserve effect&#x201d; is to be expected. For example, if this ban leads to increased productivity or redistribution of species that could benefit fisheries outside the closed area, this can offset some short-term economic losses. Thus, reducing trawling entirely in a core fishing ground may not be realistic, and anticipating effects without accounting for likely effort displacement may overestimate ecological improvements (<xref ref-type="bibr" rid="B7">Bastardie et&#xa0;al., 2014</xref>; <xref ref-type="bibr" rid="B39">Vaughan, 2017</xref>).</p>
<p>Spatial closures of trawling may be insufficient if effort is redistributed rather than reduced (<xref ref-type="bibr" rid="B5">Bastardie et&#xa0;al., 2025</xref>; <xref ref-type="bibr" rid="B8">Binch et&#xa0;al., 2025</xref>). Hence, a dynamic scenario was implemented to predict the effects of closing all MPAs in the study area and the consequent redistribution of trawling effort. This scenario is policy relevant as the Greek government is currently in the process of implementing a trawling ban in all MPAs<xref ref-type="fn" rid="fn2"><sup>2</sup></xref>. This scenario yielded minimal benthic ecosystem benefits. This is because many of the MPAs in the study area were established to meet Natura 2000 conservation objectives, i.e., the protection of habitats and species included in Natura 2000&#x2019;s annexes, rather than as fisheries management tools designed to maintain or restore seafloor integrity. Our results show that the Greek government&#x2019;s policy, while well-intentioned, may produce unintended ecological consequences if implemented without complementary management measures. Specifically, we found that MPA closures resulted in localized RBS declines in already heavily fished areas, particularly in Patraikos Gulf. In certain grid cells within this area, our model predicted RBS values falling below 0.8 - the proposed threshold for GES - as displaced fishing effort concentrated in the remaining accessible grounds. This result is concerning given that Patraikos Gulf represents one of the most economically important fishing grounds in the region, where increased effort intensity could simultaneously compromise both ecological status and long-term fishery sustainability. The displacement effect we observed complements the findings by <xref ref-type="bibr" rid="B5">Bastardie et&#xa0;al. (2025)</xref>, who showed through bioeconomic simulations that spatial restrictions can have unexpected negative outcomes when effort is reallocated to adjacent areas. Similarly, <xref ref-type="bibr" rid="B8">Binch et&#xa0;al. (2025)</xref> concluded that trawling effort displacement can transfer ecological impacts beyond protected areas and influence whole food webs in the North Sea. Despite the ecological risk, spatial restrictions and fishing effort redistribution also impose significant economic trade-offs. For example, <xref ref-type="bibr" rid="B34">Sbrana et&#xa0;al. (2025)</xref> noted from a modelling scenario, restricting fishing in shallow coastal areas (&lt;6 nm) in Italian waters, significantly affects fleet profitability by reallocating effort to other fishing grounds.</p>
<p>Overall, most habitat types in the GSA20 are currently in good condition (with RBS values higher than 0.9 in every habitat), even though some areas, particularly in circalittoral and offshore circalittoral mud habitats, exhibit higher impact from bottom trawling pressure. This is due to two reasons, firstly the Eastern Mediterranean, with the exception of some localized trawling hotspots, is not under intensive trawling pressure compared to other regions of Europe; maximum SAR in Northern European waters and the Adriatic Sea can reach values of more than 10 or 40, respectively (<xref ref-type="bibr" rid="B37">van Denderen et&#xa0;al., 2020</xref>; <xref ref-type="bibr" rid="B40">Zupa et&#xa0;al., 2025</xref>). Secondly, our benthic assessment was based on the macroinvertebrate community that tend to have lower longevity (and therefore less sensitive) than the larger sized epifaunal species that are used in other Mediterranean assessments (<xref ref-type="bibr" rid="B40">Zupa et&#xa0;al., 2025</xref>). The use of infauna-based sensitivity may be a source of underestimation of trawling impact. However, highly mobile larger epifaunal species are usually not effectively sampled by the grab devices, while trawl caught epifauna used for sensitivity estimation in other European assessments may not represent near pristine communities required for the analysis. Additionally, the spatial resolution of our study, which had C-squares of around 25 km<sup>2</sup> (resolution), may make it difficult to capture the very localized effects of bottom trawling, requiring higher resolution in habitat, sensitivity and trawling pressure maps. While this assessment is effective at characterizing impacts across broad spatial scales, they are less effective at detecting impacts on VMEs that are highly localized.</p>
<p>Together, these comparisons between the scenarios raise the question of whether large-scale spatial restrictions are necessary for seafloor integrity in areas where the baseline benthic status is already high, such as in the case of Eastern Ionian Sea. Instead, more targeted measures may be more effective, such as protection of the most sensitive habitats (e.g., maerl beds, <italic>Posidonia</italic> meadows, VMEs), reduction in overall fishing effort (which is also important for fish stocks management), gear improvements to reduce physical impact, and the use of management frameworks that minimize harmful effort displacement. These alternatives may better support the EU&#x2019;s 30 by 30 targets and Restoration Regulation objectives by focusing on ecologically valuable or vulnerable habitats.</p>
<p>Our study provides useful insights for fisheries governance in the Eastern Ionian Sea. Integrating benthic quality indicators such as RBS into spatial planning can improve alignment with EU legislation objectives, while coupling ecological assessment with socioeconomic analyses will provide a more comprehensive basis for decision-making. In this context, future research should combine RBS modeling with fleet economics, fuel consumption, carbon emissions, and target species distribution modeling at high resolution to more accurately quantify trade-offs and co-benefits, supporting sustainable fisheries management in the entire Mediterranean Sea.</p>
</sec>
<sec id="s5" sec-type="conclusions">
<label>5</label>
<title>Conclusions</title>
<p>Our study indicated that spatial management can meaningfully improve benthic status in the Eastern Ionian Sea, but the effectiveness of closures depends on the area characteristics and habitat sensitivity. Closures down to 150 m depth produced the largest improvements in benthic status, particularly in infralittoral and circalittoral habitats that host sensitive communities, but the economic costs of this type of closure might be significant since this zone has the highest concentration of bottom trawling activity (from 50 to 200 m depth). Moving the already established boundary of 1000 m maximum fishing depth to 800 or 600 m depth had negligible effects on the benthic state given there are no targeted bottom trawling activities in that zone compared to other areas in the Mediterranean Sea. In contrast, when the effort displacement triggered localized declines in the benthic status in heavily fished areas, spatial protection alone may be insufficient without complementary measures to control effort.</p>
<p>These comparisons highlighted that in regions like the Eastern Ionian Sea, where a high proportion of benthic habitats are in good status, additional ecological gains require measures that either target the most sensitive habitats (e.g. infralittoral sand, maerl beds, <italic>Posidonia</italic> meadows or vulnerable marine ecosystems) or reduce overall fishing effort. Closures at intermediate depth or minimal area restrictions are unlikely to produce measurable improvements. In addition, failure to predict displacement risks may compromise MSFD objectives and its socioeconomic benefits. Thus, future work should compare optimized management scenarios that integrate: (1) targeted protection of the most sensitive habitats or species, (2) graduated effort reduction targets coupled with spatial closures, and (3) socio-economic impact assessments to identify solutions that maximize benthic status while minimizing economic loss of fisheries.</p>
</sec>
</body>
<back>
<sec id="s6" sec-type="data-availability">
<title>Data availability statement</title>
<p>The raw data supporting the conclusions of this article will be made available by the authors, without undue reservation.</p></sec>
<sec id="s7" sec-type="author-contributions">
<title>Author contributions</title>
<p>IT: Visualization, Formal analysis, Project administration, Writing &#x2013; original draft, Data curation, Methodology, Conceptualization, Writing &#x2013; review &amp; editing. IM: Writing &#x2013; review &amp; editing, Data curation, Methodology. NP: Data curation, Writing &#x2013; review &amp; editing. FB: Methodology, Data curation, Writing &#x2013; review &amp; editing. CS: Supervision, Data curation, Writing &#x2013; review &amp; editing.</p></sec>
<ack>
<title>Acknowledgments</title>
<p>This work was supported by the EU Horizon 2020 Projects SEAwise (Shaping ecosystem-based fisheries management &#x2013; Grant Agreement No. 101000318) and GES4SEAS (Achieving Good Environmental Status for maintaining ecosystem services, by assessing integrated impacts of cumulative pressures &#x2013; Grant Agreement No. 101059877). The authors would like to acknowledge the ICES working group on Fisheries Benthic Impact and Trade-offs (WGFBIT) in development and sharing of the RBS methodology.</p>
</ack>
<sec id="s9" sec-type="COI-statement">
<title>Conflict of interest</title>
<p>The author(s) declared that this work 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="ai-statement">
<title>Generative AI statement</title>
<p>The author(s) declared that generative AI was not 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>
<ref-list>
<title>References</title>
<ref id="B1">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Agnetta</surname> <given-names>D.</given-names></name>
<name><surname>Vascotto</surname> <given-names>I.</given-names></name>
<name><surname>Panzeri</surname> <given-names>D.</given-names></name>
<name><surname>Celi&#x107;</surname> <given-names>I.</given-names></name>
<name><surname>Solidoro</surname> <given-names>C.</given-names></name>
<name><surname>Fortibuoni</surname> <given-names>T.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Bottom trawling and environmental variables drive the biodiversity of mediterranean demersal assemblages</article-title>. <source>Sci. Rep.</source> <volume>15</volume>, <fpage>27188</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1038/s41598-025-12258-2</pub-id>, PMID: <pub-id pub-id-type="pmid">40715241</pub-id>
</mixed-citation>
</ref>
<ref id="B2">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Amoroso</surname> <given-names>R. O.</given-names></name>
<name><surname>Pitcher</surname> <given-names>C. R.</given-names></name>
<name><surname>Rijnsdorp</surname> <given-names>A. D.</given-names></name>
<name><surname>McConnaughey</surname> <given-names>R. A.</given-names></name>
<name><surname>Parma</surname> <given-names>A. M.</given-names></name>
<name><surname>Suuronen</surname> <given-names>P.</given-names></name>
<etal/>
</person-group>. (<year>2018</year>). 
<article-title>Bottom trawl fishing footprints on the world&#x2019;s continental shelves</article-title>. <source>Proc. Natl. Acad. Sci.</source> <volume>115</volume>, <elocation-id>E10275-E10282</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.1802379115</pub-id>, PMID: <pub-id pub-id-type="pmid">30297399</pub-id>
</mixed-citation>
</ref>
<ref id="B3">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bahamon</surname> <given-names>N.</given-names></name>
<name><surname>Recasens</surname> <given-names>L.</given-names></name>
<name><surname>Sala-Coromina</surname> <given-names>J.</given-names></name>
<name><surname>Calero</surname> <given-names>B.</given-names></name>
<name><surname>Garcia</surname> <given-names>J. A.</given-names></name>
<name><surname>Rotllant</surname> <given-names>G.</given-names></name>
<etal/>
</person-group>. (<year>2024</year>). 
<article-title>Selectivity-based management for reversing overexploitation of demersal fisheries in North-western Mediterranean Sea</article-title>. <source>Mar. Policy</source> <volume>165</volume>, <elocation-id>106185</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.marpol.2024.106185</pub-id>
</mixed-citation>
</ref>
<ref id="B4">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
<name><surname>Angelini</surname> <given-names>S.</given-names></name>
<name><surname>Bolognini</surname> <given-names>L.</given-names></name>
<name><surname>Fuga</surname> <given-names>F.</given-names></name>
<name><surname>Manfredi</surname> <given-names>C.</given-names></name>
<name><surname>Martinelli</surname> <given-names>M.</given-names></name>
<etal/>
</person-group>. (<year>2017</year>). 
<article-title>Spatial planning for fisheries in the Northern Adriatic: Working toward viable and sustainable fishing</article-title>. <source>Ecosphere</source> <volume>8</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1002/ecs2.1696</pub-id>
</mixed-citation>
</ref>
<ref id="B5">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
<name><surname>Astarloa</surname> <given-names>A.</given-names></name>
<name><surname>Binch</surname> <given-names>L.</given-names></name>
<name><surname>Bitetto</surname> <given-names>I.</given-names></name>
<name><surname>Damalas</surname> <given-names>D.</given-names></name>
<name><surname>Depestele</surname> <given-names>J.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Anticipating how spatial fishing restrictions in EU waters perform to protect marine species, habitats, and dependent fisheries</article-title>. <source>Front. Mar. Sci.</source> <volume>12</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fmars.2025.1629180</pub-id>
</mixed-citation>
</ref>
<ref id="B6">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
<name><surname>Danto</surname> <given-names>J.</given-names></name>
<name><surname>Rufener</surname> <given-names>M. C.</given-names></name>
<name><surname>van Denderen</surname> <given-names>D.</given-names></name>
<name><surname>Eigaard</surname> <given-names>O. R.</given-names></name>
<name><surname>Dinesen</surname> <given-names>G. E.</given-names></name>
<etal/>
</person-group>. (<year>2020</year>). 
<article-title>Reducing fisheries impacts on the seafloor: A bio-economic evaluation of policy strategies for improving sustainability in the Baltic Sea</article-title>. <source>Fish Res.</source> <volume>230</volume>, <elocation-id>105681</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.fishres.2020.105681</pub-id>
</mixed-citation>
</ref>
<ref id="B7">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
<name><surname>Nielsen</surname> <given-names>J. R.</given-names></name>
<name><surname>Miethe</surname> <given-names>T.</given-names></name>
</person-group> (<year>2014</year>). 
<article-title>DISPLACE: a dynamic, individual-based model for spatial fishing planning and effort displacement &#x2014; integrating underlying fish population models</article-title>. <source>Can. J. Fisheries Aquat. Sci.</source> <volume>71</volume>, <fpage>366</fpage>&#x2013;<lpage>386</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1139/cjfas-2013-0126</pub-id>
</mixed-citation>
</ref>
<ref id="B8">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Binch</surname> <given-names>L.</given-names></name>
<name><surname>Poos</surname> <given-names>J. J.</given-names></name>
<name><surname>van de Wolfshaar</surname> <given-names>K.</given-names></name>
</person-group> (<year>2025</year>). 
<article-title>Fishing effort displacement drives ecosystem impacts within and beyond marine protected areas</article-title>. <source>Ecol. Modell</source> <volume>510</volume>, <elocation-id>111336</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/J.ECOLMODEL.2025.111336</pub-id>
</mixed-citation>
</ref>
<ref id="B9">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bona</surname> <given-names>G.</given-names></name>
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Penna</surname> <given-names>M.</given-names></name>
<name><surname>Pulcini</surname> <given-names>M.</given-names></name>
<name><surname>Raicevich</surname> <given-names>S.</given-names></name>
<name><surname>Span&#xf2;</surname> <given-names>N.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Application of a quantitative framework to estimate trawling impacts on benthic community in Central Mediterranean Sea</article-title>. <source>ICES J. Mar. Sci.</source> <volume>82</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsaf126</pub-id>
</mixed-citation>
</ref>
<ref id="B10">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Bradshaw</surname> <given-names>C.</given-names></name>
<name><surname>Iburg</surname> <given-names>S.</given-names></name>
<name><surname>Morys</surname> <given-names>C.</given-names></name>
<name><surname>Sk&#xf6;ld</surname> <given-names>M.</given-names></name>
<name><surname>Pusceddu</surname> <given-names>A.</given-names></name>
<name><surname>Ennas</surname> <given-names>C.</given-names></name>
<etal/>
</person-group>. (<year>2024</year>). 
<article-title>Effects of bottom trawling and environmental factors on benthic bacteria, meiofauna and macrofauna communities and benthic ecosystem processes</article-title>. <source>Sci. Total Environ.</source> <volume>921</volume>, <elocation-id>171076</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/J.SCITOTENV.2024.171076</pub-id>, PMID: <pub-id pub-id-type="pmid">38382611</pub-id>
</mixed-citation>
</ref>
<ref id="B11">
<mixed-citation publication-type="web">
<person-group person-group-type="author"><collab>EC</collab>
</person-group> (<year>2006</year>). 
<article-title>Council Regulation (EC) No 1967/2006 of 21 December 2006 concerning management measures for the sustainable exploitation of fishery resources in the Mediterranean Sea, and amending Regulation (EC) No 1627/94</article-title>. <source>Off. J. Eur. Union L</source> <volume>409</volume>, <fpage>11</fpage>&#x2013;<lpage>85</lpage>. Available online at: <uri xlink:href="https://eur-lex.europa.eu/legal-content/EN/TXT/?uri=CELEX%3A32006R1967">https://eur-lex.europa.eu/legal-content/EN/TXT/?uri=CELEX%3A32006R1967</uri> (<date-in-citation content-type="access-date">January 26, 2026</date-in-citation>).
</mixed-citation>
</ref>
<ref id="B12">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>EC</collab>
</person-group> (<year>2021</year>). &#x201c;
<article-title>European Commission: Directorate-General for Environment, EU biodiversity strategy for 2030 &#x2013; Bringing nature back into our lives</article-title>,&#x201d; (<publisher-loc>Luxembourg</publisher-loc>: 
<publisher-name>Office of the European Union</publisher-name>). doi:&#xa0;<pub-id pub-id-type="doi">10.2779/677548</pub-id>
</mixed-citation>
</ref>
<ref id="B13">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Eigaard</surname> <given-names>O. R.</given-names></name>
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
<name><surname>Breen</surname> <given-names>M.</given-names></name>
<name><surname>Dinesen</surname> <given-names>G. E.</given-names></name>
<name><surname>Hintzen</surname> <given-names>N. T.</given-names></name>
<name><surname>Laffargue</surname> <given-names>P.</given-names></name>
<etal/>
</person-group>. (<year>2016</year>). 
<article-title>Estimating seabed pressure from demersal trawls, seines, and dredges based on gear design and dimensions</article-title>. <source>ICES J. Mar. Sci.</source> <volume>73</volume>, <fpage>i27</fpage>&#x2013;<lpage>i43</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsv099</pub-id>
</mixed-citation>
</ref>
<ref id="B14">
<mixed-citation publication-type="web">
<person-group person-group-type="author"><collab>EU</collab>
</person-group> (<year>2024</year>). 
<article-title>Regulation (EU) 2024/1991 of The European Parliament and of the Council of 24 June 2024 on nature restoration and amending Regulation (EU) 2022/869 (Text with EEA relevance)</article-title>. Available online at: <uri xlink:href="http://data.europa.eu/eli/reg/2024/1991/oj">http://data.europa.eu/eli/reg/2024/1991/oj</uri> (Accessed <date-in-citation content-type="access-date">September 25, 2025</date-in-citation>).
</mixed-citation>
</ref>
<ref id="B15">
<mixed-citation publication-type="web">
<person-group person-group-type="author"><collab>European Commission</collab>
</person-group> (<year>2023</year>). 
<article-title>EU Action Plan: Protecting and Restoring Marine Ecosystems for Sustainable and Resilient Fisheries</article-title>. <source>Communication COM/2023/102</source>. <publisher-loc>Brussels</publisher-loc>: 
<publisher-name>European Commission</publisher-name>. Available online at: <uri xlink:href="http://oceans-and-fisheries.ec.europa.eu/policy/common-fisheries-policy-cfp/action-plan-protecting-and-restoring-marine-ecosystems-sustainable-and-resilient-fisheries_en">http://oceans-and-fisheries.ec.europa.eu/policy/common-fisheries-policy-cfp/action-plan-protecting-and-restoring-marine-ecosystems-sustainable-and-resilient-fisheries_en</uri>.
</mixed-citation>
</ref>
<ref id="B16">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>FAO</collab>
</person-group> (<year>2020</year>). &#x201c;
<article-title>The state of mediterranean and black sea fisheries 2020</article-title>,&#x201d; in <source>General fisheries commission for the mediterranean</source> (<publisher-loc>Rome</publisher-loc>: 
<publisher-name>FAO</publisher-name>). doi:&#xa0;<pub-id pub-id-type="doi">10.4060/cb2429en</pub-id>
</mixed-citation>
</ref>
<ref id="B17">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>FAO</collab>
</person-group> (<year>2021</year>). <source>GFCM 2030 Strategy for sustainable fisheries and aquaculture in the Mediterranean and the Black Sea</source> (<publisher-loc>Rome</publisher-loc>: 
<publisher-name>FAO</publisher-name>). doi:&#xa0;<pub-id pub-id-type="doi">10.4060/cb7562en</pub-id>
</mixed-citation>
</ref>
<ref id="B18">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>FAO</collab>
</person-group> (<year>2023</year>). &#x201c;
<article-title>Report of the sixteenth session of the Compliance Committee, Rhodes, Greece, 5 May 2023</article-title>,&#x201d; in <source>General Fisheries Commission for the Mediterranean/Rapport de la seizi&#xe8;me session du Comit&#xe9; d&#x2019;application, Rhodes, Gr&#xe8;ce, 5 mai 2023</source> (
<publisher-name>Commission g&#xe9;n&#xe9;rale des p&#xea;ches pour la M&#xe9;diterran&#xe9;e. FAO</publisher-name>). doi:&#xa0;<pub-id pub-id-type="doi">10.4060/cc8390b</pub-id>
</mixed-citation>
</ref>
<ref id="B19">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Fiorentino</surname> <given-names>F.</given-names></name>
<name><surname>Vitale</surname> <given-names>S.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title>How can we reduce the overexploitation of the mediterranean resources</article-title>? <source>Front. Mar. Sci.</source> <volume>8</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.3389/fmars.2021.674633</pub-id>
</mixed-citation>
</ref>
<ref id="B20">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Foden</surname> <given-names>J.</given-names></name>
<name><surname>Rogers</surname> <given-names>S.</given-names></name>
<name><surname>Jones</surname> <given-names>A.</given-names></name>
</person-group> (<year>2011</year>). 
<article-title>Human pressures on UK seabed habitats: a cumulative impact assessment</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>428</volume>, <fpage>33</fpage>&#x2013;<lpage>47</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.3354/meps09064</pub-id>
</mixed-citation>
</ref>
<ref id="B21">
<mixed-citation publication-type="book">
<person-group person-group-type="author">
<name><surname>Haynes</surname> <given-names>W.</given-names></name>
</person-group> (<year>2013</year>). &#x201c;
<article-title>Wilcoxon rank sum test</article-title>,&#x201d; in <source>Encyclopedia of systems biology</source> (
<publisher-name>Springer New York</publisher-name>, <publisher-loc>New York, NY</publisher-loc>), <fpage>2354</fpage>&#x2013;<lpage>2355</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1007/978-1-4419-9863-7_1185</pub-id>
</mixed-citation>
</ref>
<ref id="B22">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Jennings</surname> <given-names>S.</given-names></name>
<name><surname>Sciberras</surname> <given-names>M.</given-names></name>
<name><surname>Bolam</surname> <given-names>S. G.</given-names></name>
<name><surname>Cambi&#xe8;</surname> <given-names>G.</given-names></name>
<name><surname>McConnaughey</surname> <given-names>R. A.</given-names></name>
<etal/>
</person-group>. (<year>2019</year>). 
<article-title>Assessing bottom trawling impacts based on the longevity of benthic invertebrates</article-title>. <source>J. Appl. Ecol.</source> <volume>56</volume>, <fpage>1075</fpage>&#x2013;<lpage>1084</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/1365-2664.13278</pub-id>
</mixed-citation>
</ref>
<ref id="B23">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Jennings</surname> <given-names>S.</given-names></name>
<name><surname>Sciberras</surname> <given-names>M.</given-names></name>
<name><surname>Szostek</surname> <given-names>C. L.</given-names></name>
<name><surname>Hughes</surname> <given-names>K. M.</given-names></name>
<name><surname>Ellis</surname> <given-names>N.</given-names></name>
<etal/>
</person-group>. (<year>2017</year>). 
<article-title>Global analysis of depletion and recovery of seabed biota after bottom trawling disturbance</article-title>. <source>Proc. Natl. Acad. Sci. U S A</source> <volume>114</volume>, <fpage>8301</fpage>&#x2013;<lpage>8306</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.1618858114</pub-id>, PMID: <pub-id pub-id-type="pmid">28716926</pub-id>
</mixed-citation>
</ref>
<ref id="B24">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Valanko</surname> <given-names>S.</given-names></name>
<name><surname>Batts</surname> <given-names>L.</given-names></name>
<name><surname>Beukhof</surname> <given-names>E. D.</given-names></name>
<name><surname>Blomqvist</surname> <given-names>M.</given-names></name>
<name><surname>Bolam</surname> <given-names>S.</given-names></name>
<etal/>
</person-group>. (<year>2026</year>). 
<article-title>Assessment of bottom trawl impacts on the status of seabed communities in european seas</article-title>. <source>Fish Fisheries</source>. <volume>27</volume>, <fpage>285</fpage>&#x2013;<lpage>299</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/faf.70054</pub-id>
</mixed-citation>
</ref>
<ref id="B25">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Kaiser</surname> <given-names>M. J.</given-names></name>
</person-group> (<year>1998</year>). 
<article-title>Significance of bottom-fishing disturbance</article-title>. <source>Conserv. Biol.</source> <volume>12</volume>, <fpage>1230</fpage>&#x2013;<lpage>1235</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1046/j.1523-1739.1998.0120061230.x</pub-id>
</mixed-citation>
</ref>
<ref id="B26">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Maina</surname> <given-names>I.</given-names></name>
<name><surname>Kavadas</surname> <given-names>S.</given-names></name>
<name><surname>Vassilopoulou</surname> <given-names>V.</given-names></name>
<name><surname>Bastardie</surname> <given-names>F.</given-names></name>
</person-group> (<year>2021</year>). 
<article-title>Fishery spatial plans and effort displacement in the eastern Ionian Sea: A bioeconomic modelling</article-title>. <source>Ocean Coast. Manage.</source> <volume>203</volume>, <elocation-id>105456</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ocecoaman.2020.105456</pub-id>
</mixed-citation>
</ref>
<ref id="B27">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Pitcher</surname> <given-names>C. R.</given-names></name>
<name><surname>Ellis</surname> <given-names>N.</given-names></name>
<name><surname>Jennings</surname> <given-names>S.</given-names></name>
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Mazor</surname> <given-names>T.</given-names></name>
<name><surname>Kaiser</surname> <given-names>M. J.</given-names></name>
<etal/>
</person-group>. (<year>2017</year>). 
<article-title>Estimating the sustainability of towed fishing-gear impacts on seabed habitats: a simple quantitative risk assessment method applicable to data-limited fisheries</article-title>. <source>Methods Ecol. Evol.</source> <volume>8</volume>, <fpage>472</fpage>&#x2013;<lpage>480</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1111/2041-210X.12705</pub-id>
</mixed-citation>
</ref>
<ref id="B28">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Pitcher</surname> <given-names>C. R.</given-names></name>
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Jennings</surname> <given-names>S.</given-names></name>
<name><surname>Collie</surname> <given-names>J.</given-names></name>
<name><surname>Parma</surname> <given-names>A. M.</given-names></name>
<name><surname>Amoroso</surname> <given-names>R.</given-names></name>
<etal/>
</person-group>. (<year>2022</year>). 
<article-title>Trawl impacts on the relative status of biotic communities of seabed sedimentary habitats in 24 regions worldwide</article-title>. <source>Proc. Natl. Acad. Sci.</source> <volume>119</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1073/pnas.2109449119</pub-id>, PMID: <pub-id pub-id-type="pmid">34983873</pub-id>
</mixed-citation>
</ref>
<ref id="B29">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Porz</surname> <given-names>L.</given-names></name>
<name><surname>Yilmaz</surname> <given-names>R.</given-names></name>
<name><surname>Kossack</surname> <given-names>J.</given-names></name>
<name><surname>Zhang</surname> <given-names>W.</given-names></name>
<name><surname>Schrum</surname> <given-names>C.</given-names></name>
</person-group> (<year>2022</year>). 
<article-title>Modeling the impacts of bottom trawling on thesediment and particulate organic carbon distribution of the North Sea</article-title>. <fpage>13116</fpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5194/egusphere-egu22-13116</pub-id>
</mixed-citation>
</ref>
<ref id="B30">
<mixed-citation publication-type="book">
<person-group person-group-type="author"><collab>R Core Team</collab>
</person-group> (<year>2024</year>). 
<article-title>R: A language and environment for statistical computing</article-title> (<publisher-loc>Vienna, Austria</publisher-loc>: 
<publisher-name>R Foundation for Statistical Computing</publisher-name>). Available online at: <uri xlink:href="https://www.R-project.org/">https://www.R-project.org/</uri> (Accessed <date-in-citation content-type="access-date">January 16, 2026</date-in-citation>).
</mixed-citation>
</ref>
<ref id="B31">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Rees</surname> <given-names>T.</given-names></name>
</person-group> (<year>2003</year>). 
<article-title>C-Squares&#x201d;, a new spatial indexing system and its applicability to the description of Oceanographic Datasets</article-title>. <source>Oceanography</source> <volume>16</volume>, <fpage>11</fpage>&#x2013;<lpage>19</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.5670/oceanog.2003.52</pub-id>
</mixed-citation>
</ref>
<ref id="B32">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Rick</surname> <given-names>T.</given-names></name>
<name><surname>Ontiveros</surname> <given-names>M.&#xc1;.C.</given-names></name>
<name><surname>Jerardino</surname> <given-names>A.</given-names></name>
<name><surname>Mariotti</surname> <given-names>A.</given-names></name>
<name><surname>M&#xe9;ndez</surname> <given-names>C.</given-names></name>
<name><surname>Williams</surname> <given-names>A. N.</given-names></name>
</person-group> (<year>2020</year>). 
<article-title>Human-environmental interactions in Mediterranean climate regions from the Pleistocene to the Anthropocene</article-title>. <source>Anthropocene</source> <volume>31</volume>, <elocation-id>100253</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/J.ANCENE.2020.100253</pub-id>
</mixed-citation>
</ref>
<ref id="B33">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Rijnsdorp</surname> <given-names>A. D.</given-names></name>
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>van Denderen</surname> <given-names>P. D.</given-names></name>
<name><surname>Hintzen</surname> <given-names>N. T.</given-names></name>
<name><surname>Eigaard</surname> <given-names>O. R.</given-names></name>
<name><surname>Valanko</surname> <given-names>S.</given-names></name>
<etal/>
</person-group>. (<year>2020</year>). 
<article-title>Different bottom trawl fisheries have a differential impact on the status of the North Sea seafloor habitats</article-title>. <source>ICES J. Mar. Sci.</source> <volume>77</volume>, <fpage>1772</fpage>&#x2013;<lpage>1786</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsaa050</pub-id>
</mixed-citation>
</ref>
<ref id="B34">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Sbrana</surname> <given-names>A.</given-names></name>
<name><surname>Galli</surname> <given-names>S.</given-names></name>
<name><surname>Casini</surname> <given-names>M.</given-names></name>
<name><surname>Carlucci</surname> <given-names>R.</given-names></name>
<name><surname>Dassenakis</surname> <given-names>M.</given-names></name>
<name><surname>Fiorentino</surname> <given-names>F.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Exploring spatial-based management scenarios to protect the seafloor in different areas of the Mediterranean Sea</article-title>. <source>ICES J. Mar. Sci.</source> <volume>82</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsaf023</pub-id>
</mixed-citation>
</ref>
<ref id="B35">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Smith</surname> <given-names>C. J.</given-names></name>
<name><surname>Papadopoulou</surname> <given-names>N. K.</given-names></name>
<name><surname>Maina</surname> <given-names>I.</given-names></name>
<name><surname>Kavadas</surname> <given-names>S.</given-names></name>
<name><surname>Denderen</surname> <given-names>P. D.</given-names></name>
<name><surname>Katsiaras</surname> <given-names>N.</given-names></name>
<etal/>
</person-group>. (<year>2023</year>). 
<article-title>Relating benthic sensitivity and status to spatial distribution and intensity of trawling in the Eastern Mediterranean</article-title>. <source>Ecol. Indic</source> <volume>150</volume>, <elocation-id>110286</elocation-id>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/j.ecolind.2023.110286</pub-id>
</mixed-citation>
</ref>
<ref id="B36">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Tiano</surname> <given-names>J. C.</given-names></name>
<name><surname>Witbaard</surname> <given-names>R.</given-names></name>
<name><surname>Bergman</surname> <given-names>M. J. N.</given-names></name>
<name><surname>van Rijswijk</surname> <given-names>P.</given-names></name>
<name><surname>Tramper</surname> <given-names>A.</given-names></name>
<name><surname>van Oevelen</surname> <given-names>D.</given-names></name>
<etal/>
</person-group>. (<year>2019</year>). 
<article-title>Acute impacts of bottom trawl gears on benthic metabolism and nutrient cycling</article-title>. <source>ICES J. Mar. Sci.</source> <volume>76</volume>, <fpage>1917</fpage>&#x2013;<lpage>1930</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsz060</pub-id>
</mixed-citation>
</ref>
<ref id="B37">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>van Denderen</surname> <given-names>P. D.</given-names></name>
<name><surname>Bolam</surname> <given-names>S. G.</given-names></name>
<name><surname>Friedland</surname> <given-names>R.</given-names></name>
<name><surname>Hiddink</surname> <given-names>J. G.</given-names></name>
<name><surname>Nor&#xe9;n</surname> <given-names>K.</given-names></name>
<name><surname>Rijnsdorp</surname> <given-names>A. D.</given-names></name>
<etal/>
</person-group>. (<year>2020</year>). 
<article-title>Evaluating impacts of bottom trawling and hypoxia on benthic communities at the local, habitat, and regional scale using a modelling approach</article-title>. <source>ICES J. Mar. Sci.</source> <volume>77</volume>, <fpage>278</fpage>&#x2013;<lpage>289</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsz219</pub-id>
</mixed-citation>
</ref>
<ref id="B38">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Vasquez</surname> <given-names>M.</given-names></name>
<name><surname>S&#xe9;geat</surname> <given-names>B.</given-names></name>
<name><surname>Cordingley</surname> <given-names>A.</given-names></name>
<name><surname>Tilby</surname> <given-names>E.</given-names></name>
<name><surname>Wikstr&#xf6;m</surname> <given-names>S.</given-names></name>
<name><surname>Ehrnsten</surname> <given-names>E.</given-names></name>
<etal/>
</person-group> (<year>2023</year>). 
<article-title>EUSeaMap 2023, A European broad-scale seabed habitat map, Technical Report</article-title>. <source>EMODnet.</source> doi:&#xa0;<pub-id pub-id-type="doi">10.13155/97116</pub-id>
</mixed-citation>
</ref>
<ref id="B39">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Vaughan</surname> <given-names>D.</given-names></name>
</person-group> (<year>2017</year>). 
<article-title>Fishing effort displacement and the consequences of implementing Marine Protected Area management &#x2013; An English perspective</article-title>. <source>Mar. Policy</source> <volume>84</volume>, <fpage>228</fpage>&#x2013;<lpage>234</lpage>. doi:&#xa0;<pub-id pub-id-type="doi">10.1016/J.MARPOL.2017.07.007</pub-id>
</mixed-citation>
</ref>
<ref id="B40">
<mixed-citation publication-type="journal">
<person-group person-group-type="author">
<name><surname>Zupa</surname> <given-names>W.</given-names></name>
<name><surname>Carbonara</surname> <given-names>P.</given-names></name>
<name><surname>Bitetto</surname> <given-names>I.</given-names></name>
<name><surname>Casini</surname> <given-names>M.</given-names></name>
<name><surname>Maiorano</surname> <given-names>P.</given-names></name>
<name><surname>D&#x2019;Onghia</surname> <given-names>G.</given-names></name>
<etal/>
</person-group>. (<year>2025</year>). 
<article-title>Relative benthic status guides sustainable trawl management in the Adriatic-Ionian Seas</article-title>. <source>ICES J. Mar. Sci.</source> <volume>82</volume>. doi:&#xa0;<pub-id pub-id-type="doi">10.1093/icesjms/fsaf148</pub-id>
</mixed-citation>
</ref>
</ref-list>
<fn-group>
<fn id="n1" fn-type="custom" custom-type="edited-by">
<p>Edited by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/920633">Nahiduzzaman</ext-link>, WorldFish, Bangladesh</p></fn>
<fn id="n2" fn-type="custom" custom-type="reviewed-by">
<p>Reviewed by: <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/2952999">Peter J. Mitchell</ext-link>, Department of Primary Industries and Regional Development of Western Australia (DPIRD), Australia</p>
<p><ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3349534">Denise Marx</ext-link>, Leibniz Institute for Baltic Sea Research (LG), Germany</p></fn>
</fn-group>
<fn-group>
<fn id="fn1"><label>1</label>
<p><ext-link ext-link-type="uri" xlink:href="https://europe.oceana.org/press-releases/oceana-weak-implementation-eu-mediterranean-fisheries-law-leaves/">https://europe.oceana.org/press-releases/oceana-weak-implementation-eu-mediterranean-fisheries-law-leaves/</ext-link>.</p></fn>
<fn id="fn2"><label>2</label>
<p><ext-link ext-link-type="uri" xlink:href="https://www.ourocean2024.gov.gr/wp-content/uploads/2024/04/A5_OOC-9_PROPOSED-COMMITMENTS_12.4.pdf">https://www.ourocean2024.gov.gr/wp-content/uploads/2024/04/A5_OOC-9_PROPOSED-COMMITMENTS_12.4.pdf</ext-link>.</p></fn>
</fn-group>
</back>
</article>