<?xml version="1.0" encoding="UTF-8"?>
<!DOCTYPE article PUBLIC "-//NLM//DTD Journal Publishing DTD v2.3 20070202//EN" "journalpublishing.dtd">
<article article-type="research-article" dtd-version="2.3" xml:lang="EN" xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">
<front>
<journal-meta>
<journal-id journal-id-type="publisher-id">Front. Remote Sens.</journal-id>
<journal-title>Frontiers in Remote Sensing</journal-title>
<abbrev-journal-title abbrev-type="pubmed">Front. Remote Sens.</abbrev-journal-title>
<issn pub-type="epub">2673-6187</issn>
<publisher>
<publisher-name>Frontiers Media S.A.</publisher-name>
</publisher>
</journal-meta>
<article-meta>
<article-id pub-id-type="publisher-id">1650603</article-id>
<article-id pub-id-type="doi">10.3389/frsen.2025.1650603</article-id>
<article-categories>
<subj-group subj-group-type="heading">
<subject>Remote Sensing</subject>
<subj-group>
<subject>Original Research</subject>
</subj-group>
</subj-group>
</article-categories>
<title-group>
<article-title>Assessing the importance of acoustic backscatter and bottom trawling impact as predictor variables for mapping deep sea coral reefs</article-title>
<alt-title alt-title-type="left-running-head">Berry et al.</alt-title>
<alt-title alt-title-type="right-running-head">
<ext-link ext-link-type="uri" xlink:href="https://doi.org/10.3389/frsen.2025.1650603">10.3389/frsen.2025.1650603</ext-link>
</alt-title>
</title-group>
<contrib-group>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Berry</surname>
<given-names>Christopher</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/2962997/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/software/"/>
<role content-type="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/validation/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/visualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/formal-analysis/"/>
</contrib>
<contrib contrib-type="author" corresp="yes">
<name>
<surname>Lucieer</surname>
<given-names>Vanessa</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<xref ref-type="corresp" rid="c001">&#x2a;</xref>
<uri xlink:href="https://loop.frontiersin.org/people/764662/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Whittaker</surname>
<given-names>Joanne M.</given-names>
</name>
<xref ref-type="aff" rid="aff2">
<sup>2</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/243237/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/resources/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/project-administration/"/>
<role content-type="https://credit.niso.org/contributor-roles/writing-original-draft/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/supervision/"/>
<role content-type="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Althaus</surname>
<given-names>Franziska</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<uri xlink:href="https://loop.frontiersin.org/people/722144/overview"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/data-curation/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
</contrib>
<contrib contrib-type="author">
<name>
<surname>Williams</surname>
<given-names>Alan</given-names>
</name>
<xref ref-type="aff" rid="aff1">
<sup>1</sup>
</xref>
<role content-type="https://credit.niso.org/contributor-roles/Writing - review &#x26; editing/"/>
<role content-type="https://credit.niso.org/contributor-roles/methodology/"/>
<role content-type="https://credit.niso.org/contributor-roles/conceptualization/"/>
<role content-type="https://credit.niso.org/contributor-roles/investigation/"/>
<role content-type="https://credit.niso.org/contributor-roles/funding-acquisition/"/>
<role content-type="https://credit.niso.org/contributor-roles/Writing - original-draft/"/>
</contrib>
</contrib-group>
<aff id="aff1">
<sup>1</sup>
<institution>Engineering and Technology, CSIRO National Collections and Marine Infrastructure</institution>, <addr-line>Hobart</addr-line>, <addr-line>TAS</addr-line>, <country>Australia</country>
</aff>
<aff id="aff2">
<sup>2</sup>
<institution>Institute for Marine and Antarctic Studies, University of Tasmania</institution>, <addr-line>Hobart</addr-line>, <addr-line>TAS</addr-line>, <country>Australia</country>
</aff>
<author-notes>
<fn fn-type="edited-by">
<p>
<bold>Edited by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/89232/overview">Alex Cardoso Bastos</ext-link>, Federal University of Espirito Santo, Brazil</p>
</fn>
<fn fn-type="edited-by">
<p>
<bold>Reviewed by:</bold> <ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/784228/overview">Peter John Etnoyer</ext-link>, National Centers for Coastal Ocean Science (NOAA), United States</p>
<p>
<ext-link ext-link-type="uri" xlink:href="https://loop.frontiersin.org/people/3108832/overview">Khomsin Khomsin</ext-link>, Institut Teknologi Sepuluh Nopember Departemen Teknik Geomatika, Indonesia</p>
</fn>
<corresp id="c001">&#x2a;Correspondence: Christopher Berry, <email>christopher.berry@utas.edu.au</email>; Vanessa Lucieer, <email>vanessa.lucieer@utas.edu.au</email>
</corresp>
</author-notes>
<pub-date pub-type="epub">
<day>29</day>
<month>09</month>
<year>2025</year>
</pub-date>
<pub-date pub-type="collection">
<year>2025</year>
</pub-date>
<volume>6</volume>
<elocation-id>1650603</elocation-id>
<history>
<date date-type="received">
<day>20</day>
<month>06</month>
<year>2025</year>
</date>
<date date-type="accepted">
<day>04</day>
<month>09</month>
<year>2025</year>
</date>
</history>
<permissions>
<copyright-statement>Copyright &#xa9; 2025 Berry, Lucieer, Whittaker, Althaus and Williams.</copyright-statement>
<copyright-year>2025</copyright-year>
<copyright-holder>Berry, Lucieer, Whittaker, Althaus and Williams</copyright-holder>
<license xlink:href="http://creativecommons.org/licenses/by/4.0/">
<p>This is an open-access article distributed under the terms of the Creative Commons Attribution License (CC BY). The use, distribution or reproduction in other forums is permitted, provided the original author(s) and the copyright owner(s) are credited and that the original publication in this journal is cited, in accordance with accepted academic practice. No use, distribution or reproduction is permitted which does not comply with these terms.</p>
</license>
</permissions>
<abstract>
<p>Seamounts host some of the most biodiverse and vulnerable deep-sea ecosystems, as epitomised by the elevated diversity and biomass of fauna associated with extensive reefs of the frame-work forming coral <italic>Solenosmilia variabilis</italic>. In places, these vulnerable marine ecosystems (VMEs) have been substantially impacted by commercial bottom trawl fishing and face increasing threats from climate variability. Understanding both the current and historical distribution of coral reefs enables naturalness and recovery to be considered in conservation planning. Our study on the Tasmanian seamounts, an area characterised by extensive deep-sea coral reefs and historical trawling impacts, employed fine-scale Species Distribution Modelling (SDM) using acoustically derived bathymetry and backscatter data resolved to 15&#xa0;m resolution to capture fine-scale habitat heterogeneity. We used a two-model methodology to predict VME habitat distributions before and after trawling impacts across a broad region (&#x3e;480&#xa0;km<sup>2</sup>) containing multiple seamounts with peak depths ranging from 720&#x2013;2,073&#xa0;m and varied substrates, as indicated by a spectrum backscatter intensity values. Bathymetry, relative slope position, backscatter and trawling history were most influential among a suite of predictor variables. Comparison of before and after models showed a reduction in both total area and patch sizes of coral habitat consistent with areas classified as historically impacted by trawling. The reduction in extent of coral VME area was dependent on the probability threshold of habitat suitability used in the model, decreasing by 20.44%&#x2013;26.07% following trawling across a range of thresholds from 0.5&#x2013;0.85. Our study highlights the benefits to future spatially based biodiversity management initiatives that will stem from using high-resolution bathymetry and backscatter data, and information on anthropogenic impacts, in SDM predictions.</p>
</abstract>
<kwd-group>
<kwd>species distribution modelling</kwd>
<kwd>
<italic>Solenosmilia variabilis</italic>
</kwd>
<kwd>Tasmanian seamounts</kwd>
<kwd>vulnerable marine ecosystems</kwd>
<kwd>seamount</kwd>
<kwd>acoustic backscatter</kwd>
<kwd>multibeam echosounder</kwd>
</kwd-group>
<counts>
<page-count count="17"/>
</counts>
<custom-meta-wrap>
<custom-meta>
<meta-name>section-at-acceptance</meta-name>
<meta-value>Acoustic Remote Sensing</meta-value>
</custom-meta>
</custom-meta-wrap>
</article-meta>
</front>
<body>
<sec id="s1">
<title>1 Introduction</title>
<p>The deep ocean is recognised as a complex tapestry of diverse ecosystems. Among the most significant features of this underwater landscape are seamounts&#x2014;submerged mountains that rise from the seafloor forming distinct topographic features (<xref ref-type="bibr" rid="B50">Pitcher et al., 2007</xref>). Recent estimates indicate that there are over 43,000 seamounts worldwide which exceed 1,000&#xa0;m in elevation (<xref ref-type="bibr" rid="B25">Gevorgian et al., 2023</xref>). It is thought that as many as 1,000,000 smaller seamounts, from 100&#xa0;m&#x2013;1,000&#xa0;m elevation, could potentially be scattered throughout the world&#x2019;s oceans (<xref ref-type="bibr" rid="B78">Wessel, 2007</xref>). Seamounts are volcanic in origin and are often isolated features in a vast seascape (<xref ref-type="bibr" rid="B65">Staudigel and Clague, 2010</xref>). Their topography interacts with ocean currents, leading to increased productivity (<xref ref-type="bibr" rid="B79">White et al., 2007</xref>) that supports unique and often Vulnerable Marine Ecosystems (VMEs) (<xref ref-type="bibr" rid="B64">Schlacher et al., 2014</xref>).</p>
<p>VMEs are characterized by rich benthic communities, particularly deep-water corals such as <italic>S. variabilis</italic> (<xref ref-type="bibr" rid="B20">FAO, 2009</xref>). <italic>S. variabilis</italic> forms complex, three-dimensional habitats that support a myriad of communities, making it a key structural and habitat forming species (<xref ref-type="bibr" rid="B38">Koslow et al., 2001</xref>). The Tasmanian seamounts, located off the southern coast of Tasmania, Australia (<xref ref-type="fig" rid="F1">Figure 1</xref>), exemplify the role of seamounts, hosting rich ecosystems underpinned by extensive colonies of <italic>S. variabilis</italic> which form expansive reefs up to 3&#xa0;m thick at depths ranging from approximately 950 to 1,350&#xa0;m (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>).</p>
<fig id="F1" position="float">
<label>FIGURE 1</label>
<caption>
<p>Bathymetric map showing location of the Tasmanian seamounts on the continental slopes of southern Tasmania, Australia. The study area, highlighted in the blue region on Map 3, is defined by an area where high resolution multibeam echosounder (MBES) data has been acquired. The area includes a subset of the Tasmanian seamounts totaling 77 seamounts, hills, and knolls. These are predominantly located within the Huon Marine Park, which is highlighted in green. Seafloor camera transects have been acquired over a portion of the seamounts, as highlighted by the blue lines. Bathymetry data from RV <italic>Investigator</italic> voyage in 2020_v10 and <xref ref-type="bibr" rid="B23">GEBCO 2023</xref> Grid (2023). The depth range of 950&#x2013;1,350&#xa0;m, identified as the &#x2018;Zone of Coral Reef VME,&#x2019; has been previously identified (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>).</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g001.tif">
<alt-text content-type="machine-generated">Map showing the location of Huon Marine Park off the coast of Tasmania, Australia. It includes depth contours, study areas, and camera transects. Zones of Coral Reef Vulnerable Marine Ecosystems are highlighted. The map details various topographic features of the seabed, with depth marked in meters.</alt-text>
</graphic>
</fig>
<p>Despite their ecological significance, seamount ecosystems are increasingly at risk due to growing pressures from human activities and environmental changes (<xref ref-type="bibr" rid="B60">Rogers, 2019</xref>). Bottom trawl fishing, in particular, has emerged as one of the most destructive forces affecting seamount ecosystems, with documented long-lasting impacts (<xref ref-type="bibr" rid="B26">Goode et al., 2020</xref>). A stark example of this damage was observed by <xref ref-type="bibr" rid="B2">Althaus et al. (2009)</xref>, who recorded a dramatic reduction in coral cover and biodiversity on trawled Tasmanian seamounts. While the amount of coral removed from the seamounts as by-catch is unknown, nearby seamounts on the South Tasman Rise reported 1&#x2013;15 tons of coral by-catch in individual trawls during the 1997&#x2013;1998 fishing season (<xref ref-type="bibr" rid="B2">Althaus et al., 2009</xref>). Notably, <xref ref-type="bibr" rid="B80">Williams et al. (2010)</xref> found no signs of recovery even a decade after trawling had ceased, highlighting the vulnerability of these ecosystems. Compounding direct anthropogenic impacts, climate-related threats such as ocean acidification and warming further exacerbate the vulnerability of seamount ecosystems (<xref ref-type="bibr" rid="B5">Anderson et al., 2022</xref>), emphasizing the urgent need for robust conservation management strategies.</p>
<p>To develop and implement effective conservation strategies, it is crucial to understand the past, present and future distribution of seamount benthic communities (<xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>). This knowledge is essential for identifying priority areas for protection, assessing potential impacts of human activities, and designing networks of marine protected areas that can ensure the long-term preservation of these ecologically significant habitats (<xref ref-type="bibr" rid="B80">Williams et al., 2010</xref>). However, the vast expanse and inaccessibility of the deep sea present significant challenges for conducting comprehensive ecological assessments of these complex ecosystems (<xref ref-type="bibr" rid="B53">Ramirez-Llodra et al., 2011</xref>).</p>
<p>In response to these challenges, researchers are increasingly turning to predictive species distribution modelling (SDM) as a tool to map and understand seamount ecosystems (<xref ref-type="bibr" rid="B42">Lim et al., 2020</xref>). SDMs offer a cost-effective approach to predict the potential distribution of species or habitats across large, unexplored areas based on known occurrences linked to environmental variables (<xref ref-type="bibr" rid="B17">Elith and Leathwick, 2009</xref>). While these models have proven useful in conservation management (<xref ref-type="bibr" rid="B32">Guisan et al., 2013</xref>), developing accurate and reliable SDMs for deep-sea ecosystems presents numerous difficulties, the main ones being data limitations and the complexity of these environments (<xref ref-type="bibr" rid="B6">Ara&#xfa;jo et al., 2019</xref>).</p>
<p>Regional-scale SDMs, at resolutions of 500&#x2013;1,000&#xa0;m, have proven valuable for predicting the distribution of deep-sea species and habitats across large areas (<xref ref-type="bibr" rid="B31">Guinotte and Davies, 2014</xref>; <xref ref-type="bibr" rid="B4">Anderson et al., 2016b</xref>; <xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>). These models provide important insights for conservation planning and fisheries management; however, their reliability is often limited by the resolution of available environmental data (<xref ref-type="bibr" rid="B84">Winship et al., 2020</xref>) and the absence of crucial predictor variables (<xref ref-type="bibr" rid="B3">Anderson et al., 2016a</xref>). A significant ongoing challenge for regional-scale models is the lack of substrate information, which is critical for accurately predicting the distribution of benthic species. For example, <xref ref-type="bibr" rid="B4">Anderson et al. (2016b)</xref> attributed the poor performance of their regional models, as assessed through field validation (<xref ref-type="bibr" rid="B3">Anderson et al., 2016a</xref>), to a lack of substrate data. Similarly, <xref ref-type="bibr" rid="B31">Guinotte and Davies (2014)</xref> explicitly noted that their regional models likely overpredicted actual coral distribution due to the absence of data on hard substrate distribution. The importance of substrate type as a predictor of species distribution has also been repeatedly highlighted in best practice reviews of species distribution modelling (<xref ref-type="bibr" rid="B7">Bowden et al., 2021</xref>).</p>
<p>Additionally, the coarse resolution of regional models often fails to capture the fine-scale habitat heterogeneity characteristic of seamount ecosystems. This limitation can lead to significant overestimation of suitable habitat areas, as demonstrated by <xref ref-type="bibr" rid="B81">Williams et al. (2020a)</xref> in their study of Tasmanian seamounts. Their findings revealed that coral reef patches were substantially smaller (0.02&#x2013;0.16&#xa0;km<sup>2</sup>) than predicted by regional models, underscoring the need for higher-resolution approaches in areas of immediate conservation concern. Similarly, <xref ref-type="bibr" rid="B61">Rowden et al. (2017)</xref> and <xref ref-type="bibr" rid="B27">Goode et al. (2021)</xref> demonstrated the importance of fine-scale studies in understanding intra-seamount community dynamics.</p>
<p>The ability to model seamount ecosystems on a fine scale (at grid resolutions &#x3c;25&#xa0;m) has historically faced challenges similar to those encountered in regional-scale studies, particularly regarding the availability of critical predictor variables at the appropriate scale. In the case of the Tasmanian seamounts, two key factors have previously hindered the development of a comprehensive model: 1 the lack of detailed substrate information (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>) and 2 the absence of robustly quantified fishing impact data (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>).</p>
<p>In this study, we utilize a newly available high resolution acoustic backscatter data, together with the categorized fishing effort from <xref ref-type="bibr" rid="B82">Williams et al. (2020b)</xref>, to predict distributions of VME habitat formed by <italic>S. variabilis</italic> over a subset of the Tasmanian seamounts. These data present the first opportunity to map patches of reef across a broad region (&#x3e;480&#xa0;km<sup>2</sup>) at a fine scale (15&#xa0;m resolution), enabling multiple seamounts to be included in a single habitat suitability model. Our approach includes two distinct models. Model 1 predicts the current &#x2018;VME habitat&#x2019; distribution based on an image-derived observational dataset analysed against fine-scale environmental data derived from multi-beam acoustics, including backscatter and an index of bottom trawling impact as a predictor variable to all seamounts; Model 2 predicts pre-impact &#x2018;VME habitat&#x2019; based the same dataset, but excluding trawling impacts by removing both, the bottom trawling impact index predictor, and the observations on five seamounts known to have been heavily impacted by bottom trawling. This multi-model approach allows for a comparison of past and present VME habitat distribution, building upon the knowledge of past impacts to identify the potential habitats in this region (<xref ref-type="bibr" rid="B2">Althaus et al., 2009</xref>; <xref ref-type="bibr" rid="B80">Williams et al., 2010</xref>; <xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>).</p>
<p>This study builds upon the growing body of work modelling deep-water corals (<xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B27">Goode et al., 2021</xref>; <xref ref-type="bibr" rid="B5">Anderson et al., 2022</xref>; <xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>) examining the temporal changes in distribution, assessing the influence of crucial predictor variables, and evaluating the implications of these findings for conservation management. Ultimately, this work aims to assist in informing the development of management strategies that account for the dynamic nature of these ecosystems and the complex interplay of natural and anthropogenic factors influencing their distribution.</p>
</sec>
<sec sec-type="materials|methods" id="s2">
<title>2 Materials and methods</title>
<sec id="s2-1">
<title>2.1 Study area</title>
<p>The Tasmanian seamounts are located on the continental slope to the south of Tasmania, Australia (<xref ref-type="fig" rid="F1">Figure 1</xref>). They are clustered, with approximately 191 seamounts (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>) scattered over an area &#x3e;700&#xa0;km<sup>2</sup> (<xref ref-type="bibr" rid="B34">Heap and Harris, 2008</xref>). The seamount peaks range in depths from 570 to 2,400&#xa0;m and with seamount basal areas ranging from 0.2 to &#x223c;20&#xa0;km<sup>2</sup> (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>). Based on their morphology and very sparse geological sampling, the seamounts are believed to be volcanic in origin, with formative mechanisms thought to possibly relate to the Balleny Mantle Plume (<xref ref-type="bibr" rid="B18">Exon et al., 1997</xref>) or Edge Driven Convection (<xref ref-type="bibr" rid="B14">Davies and Rawlinson, 2014</xref>).</p>
<p>Our study area (<xref ref-type="fig" rid="F1">Figure 1</xref>) covers a subset of the Tasmanian seamounts, of which the majority are contained within the Huon Marine Park. The area is defined by the availability of high-resolution multibeam echosounder (MBES) data which covers approx. 486&#xa0;km<sup>2</sup> and includes 66 small seamounts with elevations from 100&#x2013;530&#xa0;m, and 7 knolls and hills with elevations from 65&#x2013;100&#xa0;m. Peak depths of these seamounts range from 720&#x2013;2,073&#xa0;m.</p>
</sec>
<sec id="s2-2">
<title>2.2 Predictor variables</title>
<p>Predictor variables describe the conditions that influence the presence of VME habitat to be modelled. For this study, most predictor variables are derived from MBES data. An additional predictor variable describes past commercial demersal trawl fishing effort.</p>
</sec>
<sec id="s2-3">
<title>2.3 Multibeam echosounder data</title>
<p>Sonar data of the seafloor in the study area were acquired during voyages IN2020_V10 and IN2021_V03 aboard Australia&#x2019;s Marine National Facility vessel, the <italic>RV Investigator,</italic> using a Kongsberg EM122, a 12&#xa0;kHz multibeam echosounder system. Line acquisition was optimized to ensure sufficient overlap between swath coverages, thus reducing the nadir artefacts in backscatter data and enhancing the overall resolution of resulting datasets (<xref ref-type="fig" rid="F2">Figure 2</xref>). The products derived from these surveys included bathymetry and backscatter. Here, bathymetry refers to the gridded product in which each cell represents depth (meters). Backscatter refers to the gridded product in which each cell represents acoustic backscatter intensity (decibels).</p>
<fig id="F2" position="float">
<label>FIGURE 2</label>
<caption>
<p>3D perspective view of the study region showing segmented multibeam sonar backscatter overlaid on bathymetry with a vertical exaggeration of 3. Backscatter denotes acoustic backscatter intensity (dB) as recorded by a MBES and is used here as a substrate proxy. Backscatter segments delineate acoustically similar patches from which the predictor &#x2018;Backscatter&#x2019; was derived.</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g002.tif">
<alt-text content-type="machine-generated">Three-dimensional landscape showing a terrain with various peaks and valleys represented in shades of gray. The shading indicates backscatter values in decibels, with a scale from 0 to negative forty-four on the left.</alt-text>
</graphic>
</fig>
<p>Bathymetry data were processed using Caris HIPS and SIPS 11.4.12. Outliers in the data were removed, and a 15&#xa0;m gridded CUBE surface was produced. This surface was imported into ArcGIS Pro v3.0.3 where Benthic Terrain Modeler and SAGA Toolbox add-ins were used to produce a suite of bathymetry-derived variables known to be useful in modelling deep-sea taxa (<xref ref-type="bibr" rid="B55">Rengstorf et al., 2012</xref>; <xref ref-type="bibr" rid="B15">Downie et al., 2021</xref>). As in previous studies, for example, <xref ref-type="bibr" rid="B61">Rowden et al. (2017)</xref>, additional variables were derived by performing focal mean analysis of the input variables to produce more generalized data, allowing for the effect of scale to be examined. This process involves using a near-neighbor approach. Neighborhood sizes of 3 &#xd7; 3, 5 &#xd7; 5, 7 &#xd7; 7 and 15 &#xd7; 15 were utilized resulting in the production of over 100 predictor variables (<xref ref-type="sec" rid="s12">Supplementary Table S1</xref>).</p>
<p>Backscatter data were processed with QPS Fledermaus Geocoder Toolbox (FMGT) 7.10.2 using beam time series and processing parameters set to flat AVG correction and a window size of 300. A mosaic with a 15&#xa0;m resolution was produced and then imported to ArcGIS Pro v3.0.3 for segmentation. Segmentation is a process where pixels with similar characteristics are grouped together (<xref ref-type="bibr" rid="B9">Che Hasan et al., 2012</xref>; <xref ref-type="bibr" rid="B48">Misiuk and Brown, 2022</xref>). A mean shift segmentation algorithm was used in ArcGIS Pro, allowing adjustments to spatial, spectral, and segment size settings to meet user needs (<xref ref-type="bibr" rid="B12">Comaniciu and Meer, 2002</xref>). A maximum spectral setting and minimal spatial setting with a segment size of 100 pixels were used. These settings maximised the apparent heterogeneity of segments relative to the mosaic&#x2019;s resolution. Segments were then used to calculate a mean backscatter value (dB) for each segment. The resulting raster was used as a substrate proxy with higher values typically indicating rock/coarse sediments and lower values finer/soft substrates (<xref ref-type="bibr" rid="B45">Lurton and Lamarche, 2015</xref>).</p>
<p>A categorical variable describing the footprints of individual seamounts was derived from bathymetry data using an approach that involved delineating the slope break from the normalized slope and normalized profile curvature (<xref ref-type="bibr" rid="B30">Grosse et al., 2009</xref>). This variable has been found useful for separating seamounts, based on variable specificity (<xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>).</p>
</sec>
<sec id="s2-4">
<title>2.4 Spatial representation of bottom fishing effort and impact</title>
<p>The study area includes seamounts exposed to commercial demersal trawling, particularly targeting orange roughy (<italic>Hoplostethus atlanticus</italic>). The trawl effort footprint overlaps the core range of <italic>S. variabilis</italic> with the most intensively trawled features showing great reductions in VME habitat (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>). The effects of this fishing are expected to be long lasting, with no observed recovery over a 10&#x2013;year period following the cessation of trawling (<xref ref-type="bibr" rid="B80">Williams et al., 2010</xref>). While the impacts of bottom trawl fishing are typically not considered in modelling deep-ocean habitats, recent studies (<xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>) have highlighted the significant impact it can have on model outputs.</p>
<p>The impacts of commercial demersal trawl fishing in the study site have previously been quantified and categorized (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>) to classify seamounts into four categories based on fishing impact on VME habitat: Heavily Impacted, Moderately Impacted, Lightly Impacted, and No Observed Impact; a fifth category of Off-Seamount was assigned to slope areas between seamount bases. The classification system used datasets including direct observation of impact (through camera surveys), locations of known trawl blocks, historical fishing data and known fishery depth range (&#x3c;1,350&#xa0;m). This classification system adopted a conservative approach to minimize the risk of false-positive classification of fishing impact. These impact classifications were appended to the &#x2018;seamount&#x2019; categorical variable, where off-seamount values were retained, and impact indexes from &#x2018;No Observed Impact&#x2019; to &#x2018;Heavily Impacted&#x2019; were appended to individual seamounts.</p>
<p>In our study area, seamounts that were not assessed as part of the previous study by <xref ref-type="bibr" rid="B82">Williams et al. (2020b)</xref>, highlighted as &#x2018;Recoded&#x2019; in <xref ref-type="fig" rid="F3">Figure 3</xref>, were classified based on proximity to fishing impacts, location on transit routes between impacted seamounts and naming of seamounts by the fishing industry.</p>
<fig id="F3" position="float">
<label>FIGURE 3</label>
<caption>
<p>Commercial demersal trawl fishing impact classifications (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>). Four categories are used to classify fishing effort on seamounts: no observed impact, low impact, moderate impact and heavy impact. Areas between seamount bases were classified as &#x2018;off-seamount.&#x2019;</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g003.tif">
<alt-text content-type="machine-generated">Map depicting Commercial demersal trawl fishing impact levels on a terrain with colored areas indicating different impacts: green for no observed impact, pink for light impact, red for moderate impact, dark red for heavy impact, black for omitted, and outlined areas for recoded sections. A scale bar indicates distance in kilometers.</alt-text>
</graphic>
</fig>
<p>Due to anomalous observations on a seamount at the western end of the site (<xref ref-type="fig" rid="F3">Figure 3</xref>, refer to &#x2018;Omitted&#x2019;), the footprint of this seamount was omitted from the fishing impact predictor variable and corresponding sonar derived predictor variables. Observations over this seamount were considered anomalous due to the high levels of VME habitat remaining, despite moderate levels of fishing effort.</p>
</sec>
<sec id="s2-5">
<title>2.5 Response variable</title>
<p>To predict the distribution of benthic taxa, a sample of locations where the taxa are present must be observed. These data, known as the response variable, informs the modelling process by indicating the taxa&#x2019;s presence relative to predictor variables. The response variable used in this study is the presence/absence of VME habitat formed by <italic>S. variabilis</italic>. This response variable was derived from seabed imagery.</p>
<p>Seabed imagery was collected during the IN2018_V06 voyage aboard Australia&#x2019;s Marine National Facility vessel, the RV <italic>Investigator</italic>, using a towed camera system. This system included a Canon EOS-1DX video camera for HD video and two Canon EOS-1DX Mark II cameras for stereo still images, viewing the seafloor at an oblique angle. The camera system was towed at a speed of 1 knot at an altitude of 2&#xa0;m (&#xb1;0.5&#xa0;m) above the seafloor, with a Sonardyne Ranger 2 USBL system enabling georeferenced logging of the cameras and associated imagery.</p>
<p>Towed camera transects were planned employing a flexible, spatially balanced design (<xref ref-type="bibr" rid="B22">Foster et al., 2020</xref>). Three transect design approaches were employed: 1 Randomized Radial, 2 Randomized Baseline, and 3 <italic>ad hoc</italic> transects conducted during transits. Each transect was typically 2&#xa0;km in length.</p>
<p>The resulting imagery was annotated using the Monterey Bay Aquarium Research Institute Video Annotation and Reference System (VARS). Reef cover was determined to represent a VME habitat by <xref ref-type="bibr" rid="B81">Williams et al. (2020a)</xref>, following a method and criterion first described by <xref ref-type="bibr" rid="B61">Rowden et al. (2017)</xref>: 25% coral reef cover with at least one live coral head in a moving window with a ratio of 3:5. This method provides a robust determination of coral cover relative to the definitions of VMEs.</p>
<p>Due to the anomalous nature of the observations on the seamount at the western end of the site (<xref ref-type="fig" rid="F3">Figure 3</xref> refer to &#x2018;Omitted&#x2019;) data from this location were omitted from the response variable. These observations were considered anomalous due to the high levels of remaining VME habitat, despite the known moderate levels of fishing impact.</p>
</sec>
<sec id="s2-6">
<title>2.6 Variable selection</title>
<p>Including redundant variables during the modeling process can lead to overfitting, which reduces the accuracy of predictions. Therefore, model performance improves by using a small number of the most relevant variables (<xref ref-type="bibr" rid="B37">Huang et al., 2011</xref>).</p>
<p>The initial suite of over 100 variables (<xref ref-type="sec" rid="s12">Supplementary Table S1</xref>), was reduced by removing variables that were highly correlated, an approach commonly used in SDMs (<xref ref-type="bibr" rid="B3">Anderson et al., 2016a</xref>; <xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B27">Goode et al., 2021</xref>). A Pearson correlation coefficient was calculated in R v4.2.2, and variable pairs/sets with r &#x3e; 0.75 were represented by a single variable, removing the rest from further analyses.</p>
<p>To further reduce the suite of variables, a preliminary Random Forest (RF) model was constructed using this reduced set of variables. Variable importance was assessed according to the mean decrease in accuracy (MSE) scaled to 100% (<xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>). This process highlighted 10 variables of the highest relevance (<xref ref-type="table" rid="T1">Table 1</xref>). These variables were further examined in an additional Pearson correlation coefficient matrix which highlighted the low correlation of the remaining variables, as demonstrated by values of r &#x3c; 0.75 (<xref ref-type="fig" rid="F4">Figure 4</xref>) except for the Seamount and Fishing Impact variable. While the Seamount and Fishing Impact variables were highly correlated, they were not both used in the same model so were retained.</p>
<table-wrap id="T1" position="float">
<label>TABLE 1</label>
<caption>
<p>The final set of variables utilized in modelling following the variable selection processes.</p>
</caption>
<table>
<thead valign="top">
<tr style="background-color:#D9D9D9">
<th align="center">Variable</th>
<th align="center">Description</th>
<th align="center">Source</th>
<th align="center">Unit</th>
<th align="center">Tool</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="left">Bathymetry</td>
<td align="left">Depth at grid cell</td>
<td align="left">MBES acquisition</td>
<td align="left">Meters</td>
<td align="left">Caris HIPS and SIPS v11.4.2</td>
</tr>
<tr>
<td align="left">Backscatter (BS)</td>
<td align="left">Mean backscatter intensity of a seafloor segment. Surrogate for substrate type</td>
<td align="left">MBES acquisition</td>
<td align="left">Decibels</td>
<td align="left">QPS FMGT v7.10.2/ArcGIS Pro v3.0.03</td>
</tr>
<tr>
<td align="left">Fine BPI</td>
<td align="left">Measure of seabed structure based on change in slope used to help identify geomorphic features</td>
<td align="left">Derived from bathymetry</td>
<td align="left">-</td>
<td align="left">ArcGIS Pro v3.0.03 (Benthic Terrain Modeler Toolbox)</td>
</tr>
<tr>
<td align="left">LS Factor (LS_Fac)</td>
<td align="left">Predicts erosion potential from slope length (L) and gradient/steepness (S) length. Focal range of 15 &#xd7; 15</td>
<td align="left">Derived from bathymetry</td>
<td align="left">Index value</td>
<td align="left">ArcGIS Pro v3.0.03 (SAGA Toolbox)</td>
</tr>
<tr>
<td align="left">Relative slope Position (Rel_Slope)</td>
<td align="left">Relative position of a grid cell on a slope. Surrogate for effects of current conditions relative to position on slope</td>
<td align="left">Derived from bathymetry</td>
<td align="left">0&#x2013;1 (Bottom-Top)</td>
<td align="left">ArcGIS Pro v3.0.03 (SAGA Toolbox)</td>
</tr>
<tr>
<td align="left">Ruggedness</td>
<td align="left">Terrain heterogeneity or complexity</td>
<td align="left">Derived from bathymetry</td>
<td align="left">0&#x2013;1 (No variation- Complete variation</td>
<td align="left">ArcGIS Pro v3.0.03 (Benthic Terrain Modeler Toolbox)</td>
</tr>
<tr>
<td align="left">Planar Curvature (Plan_Curv)</td>
<td align="left">Curvature perpendicular to the direction of maximum slope. Surrogate for convergence or divergence of flow</td>
<td align="left">Derived from bathymetry</td>
<td align="left">&#x2212;1&#x2013;1 (Laterally concave - Laterally convex)</td>
<td align="left">ArcGIS Pro v3.0.03 (Benthic Terrain Modeler Toolbox)</td>
</tr>
<tr>
<td align="left">Profile Curvature (Pro_Curve)</td>
<td align="left">Curvature parallel to direction of maximum slope. Surrogate for the acceleration and deceleration of benthic flow</td>
<td align="left">Derived from bathymetry</td>
<td align="left">0&#x2013;1 (Linear- Upwardly Concave)</td>
<td align="left">ArcGIS Pro v3.0.03 (Benthic Terrain Modeler Toolbox)</td>
</tr>
<tr>
<td align="left">Fishing Impact (Model 1 only)</td>
<td align="left">Intensity of commercial demersal trawl fishing effort appended to individual seamounts</td>
<td align="left">Sourced from <xref ref-type="bibr" rid="B82">Williams et al. (2020b)</xref>
</td>
<td align="left">Ordinal Categorical 0&#x2013;4 (0 &#x3d; Off Seamount &#x3d; 4 Heavy Effort)</td>
<td align="left">ArcGIS Pro v3.0.03</td>
</tr>
<tr>
<td align="left">Seamount (Model 2 only)</td>
<td align="left">Extents of individual seamounts</td>
<td align="left">Derived from bathymetry utilizing (<xref ref-type="bibr" rid="B30">Grosse et al., 2009</xref>) method</td>
<td align="left">Nominal Categorical 0&#x2013;1 (0 &#x3d; Off Seamount, 1 &#x3d; On Seamount)</td>
<td align="left">RStudio 2024.04.2</td>
</tr>
</tbody>
</table>
</table-wrap>
<fig id="F4" position="float">
<label>FIGURE 4</label>
<caption>
<p>Pearson correlation coefficients of the least correlated variables. Removing highly correlated variables improves model performance. Variables with a correlation coefficient of &#x3e;0.75 were not used although seamount and fishing effort which were retained and used in separate models. Variable abbreviations are given in <xref ref-type="table" rid="T1">Table 1</xref>.</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g004.tif">
<alt-text content-type="machine-generated">Correlation matrix heatmap displaying relationships among variables like Bathy, BS, Fine_BPI, Fishing, LS_Fac, Plan_Curv, Pro_Curv, Rel_Slope, Rugged, and Seamount. Color gradient from -1 to 1 indicates correlation strength, with darker colors for stronger correlations.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s2-7">
<title>2.7 Model selection and parameters</title>
<p>A Random Forest (RF) modelling approach (<xref ref-type="bibr" rid="B8">Breiman, 2001</xref>) was used to predict the spatial distribution of VME habitat formed by the coral <italic>S. variabilis</italic>.</p>
<p>RF models calculate habitat suitability using classification, or regression, trees (CART). CARTs examine the relationship between response and predictor variables by carrying out recursive binary splits of random subsets, or bootstraps, of training data (<xref ref-type="bibr" rid="B68">Strobl et al., 2009</xref>). Individual CARTs can have high variance, so ensembles of CARTs are utilized to provide a more generalized prediction (<xref ref-type="bibr" rid="B33">Hastie et al., 2009</xref>).</p>
<p>A RF approach was adopted due to its strong performance in previous studies (<xref ref-type="bibr" rid="B16">Du Preez et al., 2016</xref>; <xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>; <xref ref-type="bibr" rid="B27">Goode et al., 2021</xref>), and due to its inbuilt model performance and variable assessment metrics. The RF model was developed using the R package &#x201c;randomForest&#x201d; (<xref ref-type="bibr" rid="B41">Liaw and Wiener, 2001</xref>).</p>
<p>Two models were developed to assess the past and present extent of VME habitat across the seamounts in the study area. Model 1 incorporated all predictor variables excluding the seamount variable (nine predictor variables) and the entire response variable dataset. This model provides a conservative prediction of current probability of presence accounting for the impacts of commercial demersal trawl fishing. In Model 2 observations from seamounts classified as &#x2018;heavily impacted&#x2019; were excluded, observations from lightly and moderately impacted seamounts were retained, and the Fishing Impact predictor was removed, instead including the Seamount variable. This configuration limits the risk that disturbance-driven absences on heavily impacted seamounts are encoded as environmental limits, or false negatives, while retaining lightly and moderately impacted seamounts provides environmental contrasts under limited disturbance that help reduce false positives. Retaining these sites also maintained environmental variance and sample size for stable model estimation. Model 2 therefore provides a conservative, lower-bound VME habitat prediction of pre-impact conditions suitable for extrapolation across the region.</p>
<p>The hyperparameter &#x2018;mtry&#x2019; was tuned for each model using the tuneRF function in the randomForest package with values of 4 for both models. The optimal value for <italic>ntree</italic> was determined through establishing the rate at which errors stabilized in initial models, found to be 1,500 and 1,000 for Model 1 and 2, respectively. As classification trees are sensitive to class imbalances a down sampling approach (<xref ref-type="bibr" rid="B75">Valavi et al., 2021</xref>) was used to balance the dataset.</p>
</sec>
<sec id="s2-8">
<title>2.8 Model performance</title>
<p>Model performance was evaluated based on assessments of area under the receiver operating characteristic curve (AUC) and True Skill Statistics (TSS) which were calculated through spatial block cross-validation.</p>
<p>AUC is a threshold independent model performance metric involving the comparison of a model&#x2019;s predicted probabilities to withheld data across all possible thresholds, providing a single metric of discrimination (<xref ref-type="bibr" rid="B21">Fielding and Bell, 1997</xref>). The calculated AUC statistics range from 0&#x2013;1 with AUC values greater than 0.5 indicating better than random performance, values 0.7&#x2013;0.8 indicating adequate performance, and values greater than 0.8 indicating excellent performance (<xref ref-type="bibr" rid="B36">Hosmer et al., 2013</xref>).</p>
<p>TSS is a threshold dependent model performance metric which compares sensitivity (true prediction of presence) and specificity (correct prediction of absence). The calculated TSS statistics range from &#x2212;1 &#x2013; &#x2b;1 where &#x2212;1 indicates no better than random performance, and &#x2b;1 indicates perfect performance with values of &#x3e;0.6 considered to be useful (<xref ref-type="bibr" rid="B1">Allouche et al., 2006</xref>).</p>
<p>Performance metrics were calculated through spatial block cross-validation (<xref ref-type="bibr" rid="B74">Valavi et al., 2019</xref>). This approach accounts for spatial structuring where observations taken at geographically nearby locations are correlated (<xref ref-type="bibr" rid="B72">Tobler, 1970</xref>). Spatial structuring may result in overinflated assessment of model performance (<xref ref-type="bibr" rid="B58">Roberts et al., 2017</xref>) and is overcome by partitioning data into blocks with similar geographic locations. The block sizes used in cross-validation were calculated through assessment of the spatial autocorrelation in model residuals (<xref ref-type="bibr" rid="B69">Telford and Birks, 2009</xref>). Blocks were allocated to 1 of 10 folds, with each model being run 10 times to calculate mean AUC and TSS values.</p>
</sec>
<sec id="s2-9">
<title>2.9 Model uncertainty</title>
<p>Model uncertainty was evaluated by calculating the coefficient of variation (CV) for each cell in the prediction grid, following the methodology of <xref ref-type="bibr" rid="B24">Georgian et al. (2019)</xref> and <xref ref-type="bibr" rid="B61">Rowden et al. (2017)</xref>. The CV is a measure of relative variability, calculated as the ratio of the standard deviation to the mean. For each cell, the CV was determined using 200 model iterations. To enhance interpretability, the CV for each cell was then expressed as a percentage.</p>
<p>Converting the CV to a percentage makes it easier to understand and compare the level of uncertainty across different cells by providing a clear, standardized measure of variability relative to the mean, which can be intuitively understood as the amount of variation per 100 units of the mean value.</p>
</sec>
<sec id="s2-10">
<title>2.10 Predictor variable influence</title>
<p>The importance of each predictor variable was assessed using the <italic>randomForest</italic> package&#x2019;s inbuilt variable importance assessment. This method evaluates importance through permutation, calculating the mean decrease in accuracy (<xref ref-type="bibr" rid="B41">Liaw and Wiener, 2001</xref>). The relative influence of variables on class response was assessed utilizing partial dependence plots (PDP). PDPs indicate the average vote percentage for the class response across the range of the predictor variable, allowing for the analysis of the range for which the predictor variable contributes to class probability. The use of PDPs is particularly effective with RF models (<xref ref-type="bibr" rid="B28">Greenwell, 2017</xref>).</p>
</sec>
<sec id="s2-11">
<title>2.11 Model outputs and analysis</title>
<p>Outputs from each model included probability and uncertainty. These outputs were gridded at the resolution of the input data (15&#xa0;m grid cells). The probability outputs represent the overall probability of the presence of VME habitat. Each cell within the output gives a value from 0&#x2013;1 where 1 represents a high probability of VME habitat. The uncertainty output represents the uncertainty for each grid cell as represented by the percentage CV.</p>
<p>A comparison of predicted VME habitat area between models was made by applying a range of thresholds to probability outputs. Thresholds from 0.5&#x2013;1 were utilized to calculate areas, in km<sup>2</sup>, with the difference in areas between models providing an indication in changes in habitat extents. VME habitat patch sizes were directly compared using the threshold of 0.7 to define areas of highly probable presence (<xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>).</p>
<p>The spatial disparity in predictions from Model 1 and 2 was assessed by calculating the differences between probability outputs. A difference surface was computed by subtracting values of one probability output from another.</p>
<p>To assist in the interpretation of predictions, the mean values of each predictor variable (excluding &#x2018;Fishing Impact&#x2019; and &#x2018;Seamount&#x2019;) associated with each seamount footprint were computed.</p>
<p>These analyses were undertaken in ArcGIS Pro (v3.0.3).</p>
</sec>
</sec>
<sec sec-type="results" id="s3">
<title>3 Results</title>
<sec id="s3-1">
<title>3.1 Model performance</title>
<p>To evaluate model performance while accounting for spatial autocorrelation, spatial block cross-validation was applied (<xref ref-type="bibr" rid="B74">Valavi et al., 2019</xref>), with block sizes determined by the range of spatial autocorrelation in model residuals. The resulting block sizes were 152&#xa0;m for Model 1 and 222&#xa0;m for Model 2, equating to approximately 10 and 15 grid cells respectively at the 15&#xa0;m resolution of the environmental predictors. These relatively small block sizes indicate that spatial autocorrelation in residuals was confined to fine spatial scales, likely reflecting the influence of local environmental variation on species distributions. Model performance metrics were consistent across both models with excellent model performance, indicated by AUC values &#x3e;0.94 (<xref ref-type="bibr" rid="B36">Hosmer et al., 2013</xref>; <xref ref-type="table" rid="T2">Table 2</xref>). TSS values &#x3e;0.8 also support the usefulness of modelling outputs (<xref ref-type="bibr" rid="B1">Allouche et al., 2006</xref>). These consistently strong results align with the performance of RF models in other studies (<xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>).</p>
<table-wrap id="T2" position="float">
<label>TABLE 2</label>
<caption>
<p>Model performance metrics as assessed by spatial block cross-validation.</p>
</caption>
<table>
<thead valign="top">
<tr style="background-color:#D9D9D9">
<th align="center">Model</th>
<th align="center">Mean AUC</th>
<th align="center">Std Dev AUC</th>
<th align="center">Mean TSS</th>
<th align="center">Std Dev TSS</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="center">Model One</td>
<td align="center">0.96</td>
<td align="center">0.02</td>
<td align="center">0.84</td>
<td align="center">0.06</td>
</tr>
<tr>
<td align="center">Model Two</td>
<td align="center">0.94</td>
<td align="center">0.04</td>
<td align="center">0.81</td>
<td align="center">0.06</td>
</tr>
</tbody>
</table>
</table-wrap>
</sec>
<sec id="s3-2">
<title>3.2 Predictions of coral VME distribution</title>
<p>Strong performance by both models was also supported by a high predicted probability of coral VME habitat within the coral zone, within the seamount footprint and on seamount peaks, i.e., generally consistent with previous observations (see <xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>) (<xref ref-type="fig" rid="F5">Figures 5A,C</xref>, for Model 1 and Model 2, respectively).</p>
<fig id="F5" position="float">
<label>FIGURE 5</label>
<caption>
<p>The predicted probability of VME habitat (0%&#x2013;100%) and Uncertainty of predictions (CV (%)) on the Tasmanian seamounts within the study site for Model 1 <bold>(A, B)</bold> and Model 2 <bold>(C, D)</bold>.</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g005.tif">
<alt-text content-type="machine-generated">Map panels A, B, C, and D depict coral reef zones with varying percentage coverage and coefficient of variation (CV). A and C show percentage scales from 0 to above 0.9% in shades of blue to red, indicating coral presence. B and D display CV scales from 0 to above 30% in shades of gray to pink, denoting variation. Yellow outlines indicate the zone of coral reef Vulnerable Marine Ecosystems (VME). The scale bar marks 0 to 10 kilometers.</alt-text>
</graphic>
</fig>
<p>A map of prediction uncertainty indicated that all areas with a high probability (&#x3e;50%) of coral VME habitat presence, identified in both models, were associated with low uncertainty (&#x3c;10%) (<xref ref-type="fig" rid="F5">Figures 5B,D</xref>, for Model 1 and Model 2, respectively). In contrast, regions with relatively high uncertainty were located off seamounts and in deeper parts of the site, beyond the typical depth range of VME habitat formed by <italic>S. variabilis</italic>.</p>
<p>The most notable between-model differences in the predicted distributions of coral VME occurred on the seven heavily trawled seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamount 1&#x2013;7) where Model 2 (before trawling) predicted substantially larger suitable VME habitat areas (<xref ref-type="fig" rid="F6">Figure 6</xref>). Increased, but relatively smaller, VME habitat areas were apparent on several lightly trawled seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamount 22&#x2013;30) Between-model differences on moderately trawled seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamount 8&#x2013;21) were minimal. Unimpacted seamounts also showed little to no difference between models.</p>
<fig id="F6" position="float">
<label>FIGURE 6</label>
<caption>
<p>The difference in probability of predicted presence between Model 1 and Model 2. Seamount numbers utilized for identification purposes in analysis of results. Heavily impacted seamounts where observation data were excluded are also shown.</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g006.tif">
<alt-text content-type="machine-generated">Map depicting percentage differences in a region, with a color gradient from brown to purple indicating indicating changes between models. Zones of Coral Reef Vulnerable Marine Ecosystems (VME) are outlined in black. Areas of heavy impact, where observation data is excluded, are bordered in red. Various numbered spots are spread throughout, indicating specific locations. A scale at the bottom measures up to 10 kilometers.</alt-text>
</graphic>
</fig>
</sec>
<sec id="s3-3">
<title>3.3 Predictions of coral VME area</title>
<p>The total area of suitable VME habitat predicted by each model was compared using a range of probability thresholds from 0.5&#x2013;1. Model 1 (including trawling impact) consistently predicted smaller areas of suitable VME habitat than Model 2 (excluding trawling impact) (<xref ref-type="fig" rid="F7">Figure 7</xref>). At a threshold of 0.5, the total area of VME habitat predicted by Model 1 was 29.40&#xa0;km<sup>2</sup>, while Model 2 predicted a total area of 39.77&#xa0;km<sup>2</sup>. In both cases, the total area of suitable VME habitat declined as the threshold increased to 1.0 (probability &#x3d; certain).</p>
<fig id="F7" position="float">
<label>FIGURE 7</label>
<caption>
<p>Differences in total predicted VME habitat area based on thresholds from 0.5&#x2013;0.8 (C).</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g007.tif">
<alt-text content-type="machine-generated">Area graph showing total area in square kilometers and decrease in area percentage against varying thresholds. Model 1 is represented in pink and Model 2 in orange. A dashed line indicates percentage decrease.</alt-text>
</graphic>
</fig>
<p>A decrease in VME habitat area of 20.44%&#x2013;26.07% following trawling was predicted across the range of probability thresholds from 0.5&#x2013;0.85. However, as thresholds increased above 0.80, the difference narrowed significantly, with Model 1 predicting 14.7%&#x2013;14.3% less habitat than Model 2 for thresholds between 0.8 and 0.9. Using a probability threshold of 0.7 (<xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>), Model 1 predicted VME habitat patch sizes up to 0.91&#xa0;km<sup>2</sup> and a total area of 18.06&#xa0;km<sup>2</sup> (<xref ref-type="fig" rid="F8">Figure 8A</xref>). Model 2 predicted patch sizes up to 1.37&#xa0;km<sup>2</sup> and a total area of 22.50&#xa0;km<sup>2</sup> (<xref ref-type="fig" rid="F8">Figure 8B</xref>).</p>
<fig id="F8" position="float">
<label>FIGURE 8</label>
<caption>
<p>
<bold>(A)</bold> The predicted VME habitat patch sizes from Model 1 based on a threshold of 0.7. Seamount numbers utilized for identification purposes in analysis of results. <bold>(B)</bold> The predicted VME habitat patch sizes from Model 2 based on a threshold of 0.7. Seamount numbers utilized for identification purposes in analysis of results.</p>
</caption>
<graphic xlink:href="frsen-06-1650603-g008.tif">
<alt-text content-type="machine-generated">Two maps labeled A and B depict coral reef areas with varying color-coded zones based on area in square kilometers. The colors range from pale yellow representing the smallest areas to dark maroon for the largest. Zones not assessed are shown in gray. The maps highlight several numbered areas with boundaries marked for coral reef VME (Vulnerable Marine Ecosystems) zones. A scale bar indicates the measured distance in kilometers.</alt-text>
</graphic>
</fig>
<p>The largest predicted loss of VME habitat was observed on the heavily fished seamount 1 (<xref ref-type="fig" rid="F6">Figure 6</xref>). Model 1 predicted an almost complete loss of VME habitat on this seamount, with three significant patches being predicted as removed when using a threshold of 0.7 (<xref ref-type="fig" rid="F8">Figure 8A</xref>). On the other large, heavily fished seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamounts 2,3,6) only very small patches of VME habitat (&#x3c;0.01) were predicted with Model 1. Model 2 predicted only relatively small dispersed VME habitat patches on seamounts 2,3,6.</p>
<p>Lightly fished seamounts were consistently predicted to have decreased habitat after trawling, with seamount 22 showing the highest VME habitat loss. This seamount supported the second largest contiguous habitat patch (1.18&#xa0;km<sup>2</sup>) in Model 2 when using a threshold of 0.7. In Model 1 this large contiguous patch was reduced to over 60 small patches from 0.001&#x2013;0.09 totaling 0.43&#xa0;km<sup>2</sup>.</p>
<p>Moderately impacted seamounts exhibited minimal differences between models, with only slight decreases or increases in VME habitat area. Slight decreases in VME habitat area were predicted (&#x223c;0.05&#xa0;km<sup>2</sup> at a threshold of 0.7) on several smaller seamounts (8, 14, 20) while seamounts 10 and 21 saw no change in area. Slight increases of 0.04&#x2013;0.13&#xa0;km<sup>2</sup> (at a threshold of 0.7) were observed on seamounts 11, 12, 13, 15, 16, 17, 18, 19, 20. Seamount 9, the largest moderately fished seamount, contained the largest contiguous VME habitat patch in Model 1. In Model 2, this area decreased by 0.17&#xa0;km<sup>2</sup>. This decrease in VME habitat from Model 1 to Model 2 accounts for 0.7% of the total area and thus has minimal impact on the overall indications of changes in area. Accounting for the minor increases observed on several moderately fished seamounts, the results likely underestimate the overall reduction in coral cover caused by fishing impacts.</p>
</sec>
<sec id="s3-4">
<title>3.4 Predictor variable influence</title>
<p>Bathymetry was the most influential predictor variable in both models (<xref ref-type="table" rid="T3">Table 3</xref>), with a strong positive correlation between the 950&#x2013;1,350&#xa0;m depth range and higher coral VME habitat probability (<xref ref-type="sec" rid="s12">Supplementary Figures S1, S2</xref>). The exclusion of the fishing impact variable in Model 2 resulted in an increase in the relative importance of bathymetry, indicating its strong predictive value for VME habitat distribution. Fishing impact was highly influential in Model 1 (<xref ref-type="table" rid="T3">Table 3</xref>) and was negatively correlated with VME habitat probability in heavily fished areas. Lightly fished areas also exhibited a reduced probability of VME habitat, whereas moderately fished areas showed similar probability between models.</p>
<table-wrap id="T3" position="float">
<label>TABLE 3</label>
<caption>
<p>Predictor variable importance for Model 1 and Model 2.</p>
</caption>
<table>
<thead valign="top">
<tr style="background-color:#D9D9D9">
<th align="center">Predictor Variable</th>
<th align="center">Model 1 Importance</th>
<th align="center">Model 2 Importance</th>
</tr>
</thead>
<tbody valign="top">
<tr>
<td align="center">Bathymetry</td>
<td align="center">15.8</td>
<td align="center">26.6</td>
</tr>
<tr>
<td align="center">Fishing Impact</td>
<td align="center">15.5</td>
<td align="center">-</td>
</tr>
<tr>
<td align="center">Relative Slope Position</td>
<td align="center">14.3</td>
<td align="center">13.8</td>
</tr>
<tr>
<td align="center">Backscatter</td>
<td align="center">12.2</td>
<td align="center">13.2</td>
</tr>
<tr>
<td align="center">Planar Curvature</td>
<td align="center">9.4</td>
<td align="center">9.3</td>
</tr>
<tr>
<td align="center">Profile Curvature</td>
<td align="center">8.5</td>
<td align="center">9.1</td>
</tr>
<tr>
<td align="center">LS Factor</td>
<td align="center">8.28</td>
<td align="center">8.6</td>
</tr>
<tr>
<td align="center">Ruggedness</td>
<td align="center">8.1</td>
<td align="center">8.9</td>
</tr>
<tr>
<td align="center">Fine BPI</td>
<td align="center">7.8</td>
<td align="center">7.0</td>
</tr>
<tr>
<td align="center">Seamount</td>
<td align="center">-</td>
<td align="center">3.5</td>
</tr>
</tbody>
</table>
</table-wrap>
<p>Terrain complexity played a key role in habitat predictions. Relative slope position showed a near-linear correlation with VME habitat probability in both models (<xref ref-type="sec" rid="s12">Supplementary Figures S1, S2</xref>), with increasing values (closer to 1) indicating a preference for areas near seamount peaks. Ruggedness also contributed positively, with more structurally complex terrain associated with higher VME habitat probability. Terrain on the most heavily impacted seamounts exhibited lower mean profile curvature (i.e., flatter slopes; 0.035&#x2013;0.036, <xref ref-type="sec" rid="s12">Supplementary Figure S8</xref>) and reduced ruggedness (0.0002&#x2013;0.0009, <xref ref-type="sec" rid="s12">Supplementary Figure S10</xref>), which may reflect: 1 a loss of fine-scale structural complexity due to trawling; 2 the contribution of VME habitat itself to seafloor complexity; or 3 an inherent lack of structural complexity that made these areas more attractive to fishers. LS Factor, an index of erosion potential that increases with slope steepness and length, was positively correlated with VME habitat probability up to a value of 50. Above this threshold, habitat probability declined in Model 2 but remained stable in Model 1, possibly reflecting contrasting sensitivities to substrate stability and sediment dynamics. Curvature metrics also influenced predictions: profile curvature values around 0.05 &#x2014; indicative of convex slopes where flow is accelerating&#x2014;were associated with higher habitat probability, while planar curvature values near 0 &#x2014; representing flat or linear terrain with minimal convergent or divergent flow&#x2014;were linked to lower habitat probability. In summary, the most suitable locations for coral VME habitat were the complex, rugged, and steep terrains of seamount flanks near their peaks.</p>
<p>Backscatter emerged as a significant predictor variable in both models (<xref ref-type="table" rid="T3">Table 3</xref>). A positive correlation was observed between backscatter values approaching &#x2212;32&#xa0;dB (<xref ref-type="sec" rid="s12">Supplementary Figures S1, S2</xref>) and VME habitat probability. Probability sharply decreased as backscatter values moved away from &#x223c; &#x2212;32&#xa0;dB, indicating that this range represents an optimal substrate condition. The largest and most heavily impacted seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamounts 2, 3, 6) exhibited distinctly higher mean backscatter values (&#x2212;26.1&#xa0;dB &#xb1; 0.24) compared to other heavily impacted seamounts (&#x2212;29.5&#xa0;dB &#xb1; 3.3), moderately impacted (&#x2212;29.3&#xa0;dB &#xb1; 2.2), lightly impacted (&#x2212;28.8&#xa0;dB &#xb1; 2.7), and unimpacted areas (&#x2212;29.9&#xa0;dB &#xb1; 3.3). These patterns suggest potential differences in seafloor characteristics between heavily fished and less impacted areas.</p>
</sec>
</sec>
<sec sec-type="discussion" id="s4">
<title>4 Discussion</title>
<sec id="s4-1">
<title>4.1 The influence of predictor variables</title>
<p>The varying availability and resolution of environmental predictor variables for benthic fauna has a strong and often unknown impact on the performance and reliability of model outputs (<xref ref-type="bibr" rid="B7">Bowden et al., 2021</xref>). Those that influence the distribution of VME habitat forming deep-water corals have been well-described, although they may be unavailable, or have diminished value following data cleaning to an appropriate accuracy and/or spatial resolution when they will hinder even the most sophisticated modelling efforts (<xref ref-type="bibr" rid="B7">Bowden et al., 2021</xref>). Models that lack important environmental predictor variables have been found to provide unreliable predictions (<xref ref-type="bibr" rid="B3">Anderson et al., 2016a</xref>) and thereby limit their utility for conservation management planning. Our analysis of the Tasmanian seamounts, a relatively well-surveyed area (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>), has generated the best set of critical predictor data (such as precise backscatter measurements) to enable robust fine-scale modelling.</p>
<p>Overall, the variable that most strongly influenced the prediction of VME habitat was bathymetry; the highly correlated depth range (950&#x2013;1,350&#xa0;m) aligned tightly with the core depth ranges of <italic>S. variabilis</italic> documented at this location by several studies (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>). Our bathymetry data were more finely resolved than in historical studies (15 vs. &#x2265; 40&#xa0;m grid), and therefore better suited to the model based on video transect-derived VME distribution mapped at &#x223c;25&#xa0;m<sup>2</sup> patch resolution (&#x223c;12&#xa0;m long &#xd7; 2&#xa0;m plan view wide), and the spatial scale of variation of patches where coral VME was sparse (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>). This finer resolution was likely critical for capturing the small-scale topographic complexity of seamount flanks and peaks&#x2014;such as local ridges, ledges, and abrupt slope transitions&#x2014;which are often obscured in coarser-resolution datasets (<xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>). It also enabled the detection of discrete habitat patches across the study area, with predicted patch sizes ranging from &#x3c;0.01&#xa0;km<sup>2</sup> to a maximum of 1.37&#xa0;km<sup>2</sup>, &#x2014; exceeding the largest patch (1.16&#xa0;km<sup>2</sup>) previously reported (<xref ref-type="bibr" rid="B81">Williams et al., 2020a</xref>). Across a growing body of habitat modelling literature, bathymetry and its derivatives consistently emerge as dominant predictors of deep-sea coral and benthic community distributions (<xref ref-type="bibr" rid="B16">Du Preez et al., 2016</xref>; <xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B27">Goode et al., 2021</xref>). These variables serve as effective proxies for underlying environmental gradients, such as hydrodynamic energy, substrate type, and food availability, especially in data-limited deep-sea settings (<xref ref-type="bibr" rid="B15">Downie et al., 2021</xref>). The strong influence of bathymetry and terrain metrics in our models aligns with these previous studies and further supports the central role of fine-scale topographic structure in shaping the distribution of habitat-forming corals on seamounts.</p>
<p>Significant fishing impacts have been observed across the study site (<xref ref-type="bibr" rid="B2">Althaus et al., 2009</xref>) and have been comprehensively described, quantified, and extrapolated in a previous study (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>). In this study, fishing impact data were incorporated into Model 1, where it emerged as an influential variable. Specifically, areas classified as having &#x2018;Heavy&#x2019; fishing impact were associated with significant reductions in predicted coral presence compared to Model 2. The strong influence of the fishing impact variable aligns with recent findings (<xref ref-type="bibr" rid="B67">Stephenson et al., 2023</xref>), which highlight the critical role such data play in predicting species distributions.</p>
<p>The inclusion of trawling impact data in Model 1 resulted in consistently lower predicted VME habitat area across a range of probability thresholds. These differences were most pronounced on heavily fished seamounts, where Model 1 predicted almost complete loss of suitable VME habitat. The effect was less pronounced on lightly and moderately fished seamounts, though reductions in predicted habitat area were still observed. On moderately fished seamounts, where habitat cover differed only marginally between models, localized increases in predicted habitat were observed in some areas, despite an overall decline. This highlights the nuanced influence of fishing pressure on VME habitat where impacts may vary depending on historical fishing intensity and local environmental conditions.</p>
<p>Incorporating trawling impacts is essential for understanding coral VME distributions in this region, but quantifying fishing effects at scales comparable to high-resolution sonar-derived variables poses considerable challenges. The fishing impact data used here were at a broader spatial scale than the bathymetric and backscatter data, potentially introducing uncertainty into the model outputs (<xref ref-type="bibr" rid="B76">Vierod et al., 2014</xref>). Despite this discrepancy, <xref ref-type="bibr" rid="B51">Pittman and Brown (2011)</xref> demonstrated that integrating broad-scale variables with fine-scale data can enhance model performance, underscoring the value of combining datasets. However, the coarser resolution of the fishing impact variable in this study may limit its ability to detect fine-scale variations in fishing pressure, potentially obscuring localized areas of higher or lower impact within the broader classified zones. This limitation suggests that while the model effectively captures broad-scale habitat changes, localized impacts may require finer-resolution fishing effort data to improve predictive accuracy.</p>
<p>Substrate type is a key factor influencing the distribution of deep-water corals that form VMEs, yet it is often missing from predictive models due to a lack of high-quality, spatially consistent data (<xref ref-type="bibr" rid="B4">Anderson et al., 2016b</xref>; <xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>; <xref ref-type="bibr" rid="B5">Anderson et al., 2022</xref>). Substrate may be inferred from multibeam sonar backscatter (<xref ref-type="bibr" rid="B40">Lamarche et al., 2011</xref>), but only when acquisition follows defined standard operating procedures to ensure data quality (<xref ref-type="bibr" rid="B52">Przeslawski et al., 2019</xref>). In this study, acoustic backscatter was included in both models and had a relatively strong influence on VME predictions. A backscatter value of approximately &#x2212;32&#xa0;dB (<xref ref-type="sec" rid="s12">Supplementary Figures S1, S2</xref>) was associated with higher probabilities of VME habitat, with a sharp decline in suitability observed at values above and below this threshold.</p>
<p>In general, lower backscatter values (more negative dB) are associated with soft sediments such as mud or fine sand, while higher values (less negative) indicate harder substrates like exposed rock or consolidated reef. The peak at &#x2212;32&#xa0;dB likely reflects intermediate substrate conditions&#x2014;such as coarse gravel, mixed sediment, or encrusted surfaces&#x2014;which may offer both stability and microhabitat complexity for coral attachment. Interestingly, the highest backscatter values, which typically correspond to the hardest substrates, were not associated with the highest VME probabilities. This could appear counter-intuitive given that corals require hard substrate for attachment (<xref ref-type="bibr" rid="B56">Roberts, 2009</xref>). However, it may reflect a range of interacting factors, such as intense current scouring on bare rock, a lack of fine-scale heterogeneity, or even biological masking of the acoustic signal. Thick coral matrices themselves may attenuate or scatter the sonar return, effectively reducing the measured backscatter strength despite their presence on hard substrate beneath.</p>
<p>While backscatter thresholds appear useful for identifying suitable VME habitat, it is important to note that these values are relative; absolute interpretation would require sonar calibration (<xref ref-type="bibr" rid="B43">Lucieer and Lamarche, 2011</xref>). The importance of standardising backscatter acquisition and processing has been widely recognised. <xref ref-type="bibr" rid="B39">Lamarche and Lurton (2018)</xref> provide best-practice recommendations for improving coherence and comparability across systems. <xref ref-type="bibr" rid="B63">Schimel et al. (2018)</xref> also emphasised the need for calibration and consistent workflows to improve habitat assessments. Building on these principles, advanced acquisition and processing techniques, such as backscatter intensity normalization and oversampling, have proven effective in improving the resolution of seafloor features in deep-water environments (<xref ref-type="bibr" rid="B49">Mitchell et al., 2018</xref>). Their application using autonomous underwater vehicles (AUVs) has enabled high-resolution backscatter mapping in deep-ocean settings, helping to overcome the limitations of hull-mounted systems at depth. Incorporating similar AUV-based approaches into future VME studies could help better resolve fine-scale substrate complexity in structurally complex or rugose benthic environments, potentially improving the accuracy of predictive habitat models for VMEs in deep-ocean settings.</p>
<p>This study demonstrates how the integration of high-resolution bathymetry, backscatter data, and fishing impact information enhances our ability to model VME habitat distribution with greater accuracy. Bathymetry emerged as the strongest predictor of habitat presence, with its derivatives providing valuable proxies for environmental conditions such as water flow and current exposure (<xref ref-type="bibr" rid="B77">Walbridge et al., 2018</xref>). Backscatter data played a key role in identifying substrate characteristics, though its utility is dependent on the availability of standardized and calibrated datasets. The influence of fishing impact data was also evident, particularly in areas with heavy historical trawling, where significant habitat loss was observed. However, challenges remain in aligning fine-scale habitat models with coarser-resolution fishing data, emphasizing the need for improved spatial resolution in anthropogenic impact datasets. These findings reinforce the importance of incorporating ecologically meaningful environmental predictors into fine-scale modeling efforts (<xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B54">Ramiro-S&#xe1;nchez et al., 2019</xref>) to support deep-sea conservation and management strategies, particularly in regions where human activities continue to shape benthic ecosystems.</p>
</sec>
<sec id="s4-2">
<title>4.2 Past and present VME habitat extents</title>
<p>The distribution of coral VME habitat on the Tasmanian seamounts has been significantly impacted by historical commercial demersal trawl fishing, resulting in documented reductions in coral cover (<xref ref-type="bibr" rid="B2">Althaus et al., 2009</xref>) and seamount-scale estimates of impact severity across the regional seascape (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>). However, until now, no studies have quantified the spatial extent of VME habitat reduction due to a lack of necessary data. In this study, we provide an estimate for a sub-regional area where demersal trawling was concentrated, comparing model predictions for pre- and post-trawling conditions. This approach builds on international efforts to quantify habitat changes using predictive modeling (<xref ref-type="bibr" rid="B13">Davies and Guinotte, 2011</xref>; <xref ref-type="bibr" rid="B66">Stephenson et al., 2021</xref>; <xref ref-type="bibr" rid="B29">Gros et al., 2022</xref>).</p>
<p>Fishing impact was incorporated into Model 1, following classifications largely derived from <xref ref-type="bibr" rid="B82">Williams et al. (2020b)</xref>, while Model 2 represented pre-trawling conditions by excluding observational data from heavily fished seamounts. Observations from moderately and lightly fished sites were retained to train the model; this was not ideal but was necessary because very few seamounts had experienced zero historical trawling. This therefore results in a conservative historical estimate of pre-trawling coral VME distribution by likely underestimating its extent in some areas where trawling impacts were not recognised.</p>
<p>Heavily trawled seamounts exhibited the most substantial declines in VME habitat, with Model 1 predicting near-complete loss on the largest impacted seamounts (<xref ref-type="fig" rid="F6">Figure 6</xref>: seamounts 1&#x2013;7). This is consistent with global patterns, where historically trawled seamounts have exhibited extensive habitat degradation (<xref ref-type="bibr" rid="B10">Clark et al., 2015</xref>; <xref ref-type="bibr" rid="B59">Rogers, 2018</xref>). On these heavily fished seamounts, Model 2 predicted only small patches of suitable habitat (&#x3c;0.14&#xa0;km<sup>2</sup>) near seamount peaks and flanks prior to fishing. The absence of coral rubble in previous seafloor observations at these sites further supports this result (<xref ref-type="bibr" rid="B82">Williams et al., 2020b</xref>) suggesting that environmental conditions, such as reduced terrain complexity, lower profile curvature, and differing substrate (indicated by higher backscatter values) may have been less favorable for the formation and persistence of VME habitat.</p>
<p>VME habitat fragmentation, as a result of trawling, was evident across the study area, with the most pronounced effects observed on the largest lightly fished seamount, seamount 22 (<xref ref-type="fig" rid="F6">Figure 6</xref>). This seamount exhibited the greatest degree of fragmentation, where a contiguous patch of 1.18&#xa0;km<sup>2</sup> in Model 2 (pre-trawling) was reduced to over 60 small patches (&#x3c;0.09&#xa0;km<sup>2</sup> each) in Model 1 (post-trawling). Model predictions suggest that even low levels of fishing activity can lead to significant habitat fragmentation, potentially reducing connectivity between coral communities. Such fragmentation has been identified as a factor limiting recovery potential for deep-sea coral ecosystems following trawling disturbance (<xref ref-type="bibr" rid="B26">Goode et al., 2020</xref>).</p>
<p>In contrast, seamounts classified as &#x2018;Moderately&#x2019; impacted exhibited relatively high predicted VME habitat probabilities, often exceeding those of lightly fished or unimpacted seamounts. This likely reflects their initially high coral cover, which may have made them attractive targets for trawling activity. Interestingly, Model 2 (representing pre-trawling conditions) predicted slightly higher habitat coverage on moderately fished seamounts than Model 1 (post-trawling). While this might appear counterintuitive given their classification, it is likely a result of residual influence from high initial coral abundance in the model training data, which continued to drive predictions in both models. Thus, the apparent slight increase is not interpreted as a genuine expansion of habitat post-disturbance, but more likely an artefact of how remaining presence data influenced predictions. These results suggest that moderately impacted seamounts, despite experiencing some habitat loss, may still retain more suitable habitat than lightly fished seamounts, which were potentially less favorable for coral VME development even prior to trawling.</p>
<p>These results highlight the value of high-resolution habitat modeling for improving our understanding of the distribution and spatial configuration of coral VMEs on the Tasmanian seamounts. The comparison between pre- and post-trawling scenarios illustrates how fine-scale predictions can reveal shifts in habitat extent and fragmentation that may not be detectable at broader scales. The ability to identify small, spatially distinct patches of suitable habitat&#x2014;often located on seamount flanks or ridge features&#x2014;demonstrates the benefits of incorporating detailed bathymetric, substrate, and impact data into modeling efforts. These insights provide a stronger basis for spatial planning and resource management in complex deep-sea environments, and contribute to broader efforts to assess habitat change, resilience, and recovery potential over time.</p>
</sec>
<sec id="s4-3">
<title>4.3 Implications for environmental management</title>
<p>VME habitat forming deep-sea corals are highly vulnerable to anthropogenic disturbances, including bottom trawling (<xref ref-type="bibr" rid="B11">Clark et al., 2019</xref>), mining (<xref ref-type="bibr" rid="B64">Schlacher et al., 2014</xref>), and climate change (<xref ref-type="bibr" rid="B57">Roberts and Cairns, 2014</xref>). Given their slow growth rates, with reefs taking centuries or even millennia to develop (<xref ref-type="bibr" rid="B19">Fallon et al., 2014</xref>), these ecosystems require proactive and precautionary management approaches. In this context, fine-scale, spatially explicit habitat modeling offers a valuable tool for identifying ecologically significant or sensitive areas and supporting informed conservation efforts.</p>
<p>Broad-scale and high-resolution modeling approaches are particularly suited to informing detailed reserve design, especially at local and sub-regional scales where fine-scale habitat heterogeneity is most relevant (<xref ref-type="bibr" rid="B29">Gros et al., 2022</xref>). Models at this resolution align with global best-practice recommendations&#x2014;including those of the Food and Agriculture Organization (<xref ref-type="bibr" rid="B20">FAO, 2009</xref>) &#x2014;which emphasize the importance of applying the best available scientific information, including predictive models, to guide spatial closures and mitigation strategies. These approaches are also consistent with broader international frameworks such as the Convention on Biological Diversity and its Kunming-Montreal Global Biodiversity Framework, which targets the protection of 30% of marine areas by 2030.</p>
<p>In our study, the differences observed between the pre- and post-trawling model predictions highlight how historical demersal fishing activity may have influenced the present-day distribution of suitable coral habitat, a pattern also reported in other deep-sea regions (<xref ref-type="bibr" rid="B11">Clark et al., 2019</xref>; <xref ref-type="bibr" rid="B26">Goode et al., 2020</xref>). Within the study area, several seamounts showed reduced suitability in the post-trawling model, reinforcing the importance of integrating historical disturbance data into predictive frameworks when evaluating current habitat conditions and setting conservation priorities.</p>
<p>Importantly, the modeling framework developed in this study is scalable and adaptable. It provides a practical foundation for application in other data-limited deep-sea regions, particularly where multibeam bathymetry and other environmental datasets are available. As global initiatives such as the UN Decade of Ocean Science for Sustainable Development (<xref ref-type="bibr" rid="B62">Ryabinin et al., 2019</xref>) seek to enhance ocean knowledge and inform marine spatial planning, tools that integrate acoustic, ecological, and anthropogenic impact data at high spatial resolution will become increasingly relevant in helping to support adaptive and informed management responses.</p>
<p>Looking ahead, the continued development of high-resolution, spatially explicit models will be essential as deep-sea ecosystems face mounting pressures from industrial activity and climate-related stressors (<xref ref-type="bibr" rid="B53">Ramirez-Llodra et al., 2011</xref>). These models can enhance our ability to assess habitat connectivity, identify potential recovery areas, and refine conservation strategies in line with changing ocean conditions. VME-forming deep-sea corals act as biodiversity hotspots, supporting diverse benthic communities (<xref ref-type="bibr" rid="B35">Henry and Roberts, 2017</xref>; <xref ref-type="bibr" rid="B83">Williams et al., 2021</xref>; <xref ref-type="bibr" rid="B46">Maguire et al., 2023</xref>). Their persistence may also bolster the broader resilience of marine ecosystems (<xref ref-type="bibr" rid="B70">Thurber et al., 2014</xref>), particularly as species seek climate refugia in the face of ocean warming and acidification (<xref ref-type="bibr" rid="B71">Tittensor et al., 2010</xref>). Protecting these habitats through science-based, data-driven approaches will be critical to ensuring their long-term viability.</p>
</sec>
</sec>
<sec sec-type="conclusion" id="s5">
<title>5 Conclusion</title>
<p>Modelling is increasingly being used to predict and understand the spatial patterns of biodiversity, particularly in the deep sea (<xref ref-type="bibr" rid="B61">Rowden et al., 2017</xref>; <xref ref-type="bibr" rid="B24">Georgian et al., 2019</xref>; <xref ref-type="bibr" rid="B27">Goode et al., 2021</xref>). However, SDMs still face significant challenges due to the lack of spatially explicit data, including fine-scale environmental data, data quantifying anthropogenic impacts, and the uncertainty in the temporal methods to predict species distributions over time. Despite these obstacles, our study highlights the utility of fine-scale modelling in capturing critical habitat features, such as VME habitat forming deep-water coral reef patches, and demonstrates the importance of incorporating both substrate information and fishing impacts into SDM predictions.</p>
<p>Our findings, which predict a reduction in VME habitat due to commercial demersal trawl fishing, emphasize the need to account for human impacts in biodiversity models. Although this estimate is likely conservative, and the actual loss of VME habitat may be higher, it offers a valuable baseline for guiding future management strategies. It also highlights the urgent need for more comprehensive assessments of how human activities affect deep-sea ecosystems.</p>
<p>To improve our understanding of deep-ocean ecosystems, high-resolution acoustic multibeam bathymetric and backscatter data and correlated seafloor imagery are critically needed. Initiatives like Seabed 2030 (<xref ref-type="bibr" rid="B47">Mayer et al., 2018</xref>), Seamap Australia (<xref ref-type="bibr" rid="B44">Lucieer et al., 2019</xref>) and AusSeabed (<xref ref-type="bibr" rid="B73">Townsend et al., 2023</xref>) are advancing efforts to provide accessible, high-quality seafloor mapping data adhering to FAIR data standards and incorporating internationally consistent classification models. Expanding on these efforts is essential, particularly through open access to substrate data, which is crucial for refining SDMs and informing more effective marine conservation strategies.</p>
</sec>
</body>
<back>
<sec sec-type="data-availability" id="s6">
<title>Data availability statement</title>
<p>The datasets presented in this study can be found in online repositories. The names of the repository/repositories and accession number(s) can be found in the article/<ext-link ext-link-type="uri" xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="https://doi.org/10.25919/knqv-2t21">https://doi.org/10.25919/knqv-2t21</ext-link>.</p>
</sec>
<sec sec-type="author-contributions" id="s7">
<title>Author contributions</title>
<p>CB: Investigation, Software, Writing &#x2013; review and editing, Resources, Validation, Writing &#x2013; original draft, Data curation, Project administration, Methodology, Visualization, Formal Analysis. VL: Conceptualization, Writing &#x2013; original draft, Funding acquisition, Writing &#x2013; review and editing, Resources, Methodology, Project administration, Supervision. JW: Resources, Funding acquisition, Project administration, Writing &#x2013; original draft, Conceptualization, Supervision, Writing &#x2013; review and editing. FA: Methodology, Writing &#x2013; review and editing, Data curation, Conceptualization. AW: Writing &#x2013; review and editing, Methodology, Conceptualization, Investigation, Funding acquisition, Writing &#x2013; original draft.</p>
</sec>
<sec sec-type="funding-information" id="s8">
<title>Funding</title>
<p>The author(s) declare that financial support was received for the research and/or publication of this article. Project data were collected during a voyage on Australia&#x2019;s Marine National Facility Vessel, RV <italic>Investigator</italic>, funded through the Australian Government. Modelling, data analysis and writing were partially funded by CSIRO.</p>
</sec>
<ack>
<p>The authors wish to thank and acknowledge the support staff and marine crew aboard CSIRO&#x2019;s RV <italic>Investigator</italic> (<ext-link ext-link-type="uri" xlink:href="https://ror.org/01mae9353">https://ror.org/01mae9353</ext-link>) for facilitating the data acquisition associated with this project. We also thank and acknowledge the many other engineering, technical and administrative staff at CSIRO who contributed to supporting the project that provided the data for this manuscript. We thank and acknowledge input and guidance on modelling from Nicole Hill, Skip Woolley and Jan Jansen.</p>
</ack>
<sec sec-type="COI-statement" id="s9">
<title>Conflict of interest</title>
<p>The authors declare that the research was conducted in the absence of any commercial or financial relationships that could be construed as a potential conflict of interest.</p>
</sec>
<sec sec-type="ai-statement" id="s10">
<title>Generative AI statement</title>
<p>The author(s) declare that no Generative AI was used in the creation of this manuscript.</p>
<p>Any alternative text (alt text) provided alongside figures in this article has been generated by Frontiers with the support of artificial intelligence and reasonable efforts have been made to ensure accuracy, including review by the authors wherever possible. If you identify any issues, please contact us.</p>
</sec>
<sec sec-type="disclaimer" id="s11">
<title>Publisher&#x2019;s note</title>
<p>All claims expressed in this article are solely those of the authors and do not necessarily represent those of their affiliated organizations, or those of the publisher, the editors and the reviewers. Any product that may be evaluated in this article, or claim that may be made by its manufacturer, is not guaranteed or endorsed by the publisher.</p>
</sec>
<sec sec-type="supplementary-material" id="s12">
<title>Supplementary material</title>
<p>The Supplementary Material for this article can be found online at: <ext-link ext-link-type="uri" xlink:href="https://www.frontiersin.org/articles/10.3389/frsen.2025.1650603/full#supplementary-material">https://www.frontiersin.org/articles/10.3389/frsen.2025.1650603/full&#x23;supplementary-material</ext-link>
</p>
<supplementary-material xlink:href="DataSheet1.docx" id="SM1" mimetype="application/docx" xmlns:xlink="http://www.w3.org/1999/xlink"/>
</sec>
<ref-list>
<title>References</title>
<ref id="B1">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Allouche</surname>
<given-names>O.</given-names>
</name>
<name>
<surname>Tsoar</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Kadmon</surname>
<given-names>R.</given-names>
</name>
</person-group> (<year>2006</year>). <article-title>Assessing the accuracy of species distribution models: prevalence, kappa and the true skill statistic (TSS)</article-title>. <source>J. Appl. Ecol.</source> <volume>43</volume> (<issue>6</issue>), <fpage>1223</fpage>&#x2013;<lpage>1232</lpage>. <pub-id pub-id-type="doi">10.1111/j.1365-2664.2006.01214.x</pub-id>
</citation>
</ref>
<ref id="B2">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Schlacher</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Kloser</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Green</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Barker</surname>
<given-names>B.</given-names>
</name>
<etal/>
</person-group> (<year>2009</year>). <article-title>Impacts of bottom trawling on deep-coral ecosystems of seamounts are long-lasting</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>397</volume>, <fpage>279</fpage>&#x2013;<lpage>294</lpage>. <pub-id pub-id-type="doi">10.3354/meps08248</pub-id>
</citation>
</ref>
<ref id="B3">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Guinotte</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Mormede</surname>
<given-names>S.</given-names>
</name>
<name>
<surname>Davies</surname>
<given-names>A. J.</given-names>
</name>
<etal/>
</person-group> (<year>2016a</year>). <article-title>Field validation of habitat suitability models for vulnerable marine ecosystems in the South Pacific Ocean: implications for the use of broad-scale models in fisheries management</article-title>. <source>Ocean and Coast. Manag.</source> <volume>120</volume>, <fpage>110</fpage>&#x2013;<lpage>126</lpage>. <pub-id pub-id-type="doi">10.1016/j.ocecoaman.2015.11.025</pub-id>
</citation>
</ref>
<ref id="B4">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Guinotte</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Tracey</surname>
<given-names>D. M.</given-names>
</name>
<name>
<surname>Mackay</surname>
<given-names>K. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
</person-group> (<year>2016b</year>). <article-title>Habitat suitability models for predicting the occurrence of vulnerable marine ecosystems in the seas around New Zealand</article-title>. <source>Deep Sea Res. Part I Oceanogr. Res. Pap.</source> <volume>115</volume>, <fpage>265</fpage>&#x2013;<lpage>292</lpage>. <pub-id pub-id-type="doi">10.1016/j.dsr.2016.07.006</pub-id>
</citation>
</ref>
<ref id="B5">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Stephenson</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Behrens</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Predicting the effects of climate change on deep&#x2010;water coral distribution around New Zealand&#x2014;will there be suitable refuges for protection at the end of the 21st century?</article-title> <source>Glob. Change Biol.</source> <volume>28</volume> (<issue>22</issue>), <fpage>6556</fpage>&#x2013;<lpage>6576</lpage>. <pub-id pub-id-type="doi">10.1111/gcb.16389</pub-id>
<pub-id pub-id-type="pmid">36045501</pub-id>
</citation>
</ref>
<ref id="B6">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ara&#xfa;jo</surname>
<given-names>M. B.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>R. P.</given-names>
</name>
<name>
<surname>M&#xe1;rcia Barbosa</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Beale</surname>
<given-names>C. M.</given-names>
</name>
<name>
<surname>Dormann</surname>
<given-names>C. F.</given-names>
</name>
<name>
<surname>Early</surname>
<given-names>R.</given-names>
</name>
<etal/>
</person-group> (<year>2019</year>). <article-title>Standards for distribution models in biodiversity assessments</article-title>. <source>Sci. Adv.</source> <volume>5</volume> (<issue>1</issue>), <fpage>eaat4858</fpage>. <pub-id pub-id-type="doi">10.1126/sciadv.aat4858</pub-id>
<pub-id pub-id-type="pmid">30746437</pub-id>
</citation>
</ref>
<ref id="B7">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Stephenson</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Assessing habitat suitability models for the deep sea: is our ability to predict the distributions of seafloor fauna improving?</article-title> <source>Front. Mar. Sci.</source> <volume>8</volume>, <fpage>632389</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2021.632389</pub-id>
</citation>
</ref>
<ref id="B8">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Breiman</surname>
<given-names>L.</given-names>
</name>
</person-group> (<year>2001</year>). <article-title>Random forests</article-title>. <source>Mach. Learn.</source> <volume>45</volume> (<issue>1</issue>), <fpage>5</fpage>&#x2013;<lpage>32</lpage>. <pub-id pub-id-type="doi">10.1023/a:1010933404324</pub-id>
</citation>
</ref>
<ref id="B9">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Che Hasan</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Ierodiaconou</surname>
<given-names>D.</given-names>
</name>
<name>
<surname>Laurenson</surname>
<given-names>L.</given-names>
</name>
</person-group> (<year>2012</year>). <article-title>Combining angular response classification and backscatter imagery segmentation for benthic biological habitat mapping</article-title>. <source>Estuar. Coast. Shelf Sci.</source> <volume>97</volume>, <fpage>1</fpage>&#x2013;<lpage>9</lpage>. <pub-id pub-id-type="doi">10.1016/j.ecss.2011.10.004</pub-id>
</citation>
</ref>
<ref id="B10">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Schlacher</surname>
<given-names>T. A.</given-names>
</name>
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>The impacts of deep-sea fisheries on benthic communities: a review</article-title>. <source>ICES J. Mar. Sci.</source> <volume>73</volume> (<issue>Suppl. l_1</issue>), <fpage>i51</fpage>&#x2013;<lpage>i69</lpage>. <pub-id pub-id-type="doi">10.1093/icesjms/fsv123</pub-id>
</citation>
</ref>
<ref id="B11">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Stewart</surname>
<given-names>R.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Little evidence of benthic community resilience to bottom trawling on seamounts after 15 years</article-title>. <source>Front. Mar. Sci.</source> <volume>6</volume>, <fpage>63</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2019.00063</pub-id>
</citation>
</ref>
<ref id="B12">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Comaniciu</surname>
<given-names>D.</given-names>
</name>
<name>
<surname>Meer</surname>
<given-names>P.</given-names>
</name>
</person-group> (<year>2002</year>). <article-title>Mean shift: a robust approach toward feature space analysis</article-title>. <source>IEEE Trans. Pattern Analysis Mach. Intell.</source> <volume>24</volume> (<issue>5</issue>), <fpage>603</fpage>&#x2013;<lpage>619</lpage>. <pub-id pub-id-type="doi">10.1109/34.1000236</pub-id>
</citation>
</ref>
<ref id="B13">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Davies</surname>
<given-names>A. J.</given-names>
</name>
<name>
<surname>Guinotte</surname>
<given-names>J. M.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Global habitat suitability for framework-forming cold-water corals</article-title>. <source>PLoS ONE</source> <volume>6</volume> (<issue>4</issue>), <fpage>e18483</fpage>. <pub-id pub-id-type="doi">10.1371/journal.pone.0018483</pub-id>
<pub-id pub-id-type="pmid">21525990</pub-id>
</citation>
</ref>
<ref id="B14">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Davies</surname>
<given-names>D. R.</given-names>
</name>
<name>
<surname>Rawlinson</surname>
<given-names>N.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>On the origin of recent intraplate volcanism in Australia</article-title>. <source>Geology</source> <volume>42</volume> (<issue>12</issue>), <fpage>1031</fpage>&#x2013;<lpage>1034</lpage>. <pub-id pub-id-type="doi">10.1130/g36093.1</pub-id>
</citation>
</ref>
<ref id="B15">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Downie</surname>
<given-names>A.-L.</given-names>
</name>
<name>
<surname>Vieira</surname>
<given-names>R. P.</given-names>
</name>
<name>
<surname>Hogg</surname>
<given-names>O. T.</given-names>
</name>
<name>
<surname>Darby</surname>
<given-names>C.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Distribution of vulnerable marine ecosystems at the South sandwich islands: results from the blue belt discovery expedition 99 deep-water camera surveys</article-title>. <source>Front. Mar. Sci.</source> <volume>8</volume>, <fpage>662285</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2021.662285</pub-id>
</citation>
</ref>
<ref id="B16">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Du Preez</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Curtis</surname>
<given-names>J. M. R.</given-names>
</name>
<name>
<surname>Clarke</surname>
<given-names>M. E.</given-names>
</name>
</person-group> (<year>2016</year>). <article-title>The structure and distribution of benthic communities on a shallow seamount (cobb seamount, northeast pacific ocean)</article-title>. <source>PLOS ONE</source> <volume>11</volume> (<issue>10</issue>), <fpage>e0165513</fpage>. <pub-id pub-id-type="doi">10.1371/journal.pone.0165513</pub-id>
<pub-id pub-id-type="pmid">27792782</pub-id>
</citation>
</ref>
<ref id="B17">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Elith</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Leathwick</surname>
<given-names>J.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>Species distribution models: ecological explanation and prediction across space and time</article-title>. <source>Annu. Rev. Ecol. Evol. Syst.</source> <volume>40</volume>, <fpage>677</fpage>&#x2013;<lpage>697</lpage>. <pub-id pub-id-type="doi">10.1146/annurev.ecolsys.110308.120159</pub-id>
</citation>
</ref>
<ref id="B18">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Exon</surname>
<given-names>N. F.</given-names>
</name>
<name>
<surname>Berry</surname>
<given-names>R. F.</given-names>
</name>
<name>
<surname>Crawford</surname>
<given-names>A. J.</given-names>
</name>
<name>
<surname>Hill</surname>
<given-names>P. J.</given-names>
</name>
</person-group> (<year>1997</year>). <article-title>Geological evolution of the east tasman plateau, a continental fragment southeast of Tasmania</article-title>. <source>Aust. J. Earth Sci.</source> <volume>44</volume> (<issue>5</issue>), <fpage>597</fpage>&#x2013;<lpage>608</lpage>. <pub-id pub-id-type="doi">10.1080/08120099708728339</pub-id>
</citation>
</ref>
<ref id="B19">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Fallon</surname>
<given-names>S. J.</given-names>
</name>
<name>
<surname>Thresher</surname>
<given-names>R. E.</given-names>
</name>
<name>
<surname>Adkins</surname>
<given-names>J.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Age and growth of the cold-water scleractinian Solenosmilia variabilis and its reef on SW Pacific seamounts</article-title>. <source>Coral Reefs</source> <volume>33</volume> (<issue>1</issue>), <fpage>31</fpage>&#x2013;<lpage>38</lpage>. <pub-id pub-id-type="doi">10.1007/s00338-013-1097-y</pub-id>
</citation>
</ref>
<ref id="B20">
<citation citation-type="book">
<collab>FAO</collab> (<year>2009</year>). <source>International guidelines for the management of deep-sea fisheries in the high-seas</source>. <publisher-loc>Rome</publisher-loc>: <publisher-name>Food and Agriculture Organisation of the United Nations</publisher-name>.</citation>
</ref>
<ref id="B21">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Fielding</surname>
<given-names>A. H.</given-names>
</name>
<name>
<surname>Bell</surname>
<given-names>J. F.</given-names>
</name>
</person-group> (<year>1997</year>). <article-title>A review of methods for the assessment of prediction errors in conservation presence/absence models</article-title>. <source>Environ. Conserv.</source> <volume>24</volume> (<issue>1</issue>), <fpage>38</fpage>&#x2013;<lpage>49</lpage>. <pub-id pub-id-type="doi">10.1017/s0376892997000088</pub-id>
</citation>
</ref>
<ref id="B22">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Foster</surname>
<given-names>S. D.</given-names>
</name>
<name>
<surname>Hosack</surname>
<given-names>G. R.</given-names>
</name>
<name>
<surname>Monk</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Lawrence</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Barrett</surname>
<given-names>N. S.</given-names>
</name>
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<etal/>
</person-group> (<year>2020</year>). <article-title>Spatially balanced designs for transect&#x2010;based surveys</article-title>. <source>Methods Ecol. Evol.</source> <volume>11</volume> (<issue>1</issue>), <fpage>95</fpage>&#x2013;<lpage>105</lpage>. <pub-id pub-id-type="doi">10.1111/2041-210x.13321</pub-id>
</citation>
</ref>
<ref id="B23">
<citation citation-type="journal">
<collab>GEBCO</collab> (<year>2023</year>). <article-title>GEBCO compilation group</article-title>
<italic>.</italic> <comment>Available online at:</comment> https://www.gebco.net/data-products/gridded-bathymetry-data/gebco2023-grid <pub-id pub-id-type="doi">10.5285/f98b053b-0cbc-6c23-e053-6c86abc0af7b</pub-id>
</citation>
</ref>
<ref id="B24">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Georgian</surname>
<given-names>S. E.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>Ensemble habitat suitability modeling of vulnerable marine ecosystem indicator taxa to inform deep-sea fisheries management in the South Pacific Ocean</article-title>. <source>Fish. Res.</source> <volume>211</volume>, <fpage>256</fpage>&#x2013;<lpage>274</lpage>. <pub-id pub-id-type="doi">10.1016/j.fishres.2018.11.020</pub-id>
</citation>
</ref>
<ref id="B25">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gevorgian</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Sandwell</surname>
<given-names>D. T.</given-names>
</name>
<name>
<surname>Yu</surname>
<given-names>Y.</given-names>
</name>
<name>
<surname>Kim</surname>
<given-names>S. S.</given-names>
</name>
<name>
<surname>Wessel</surname>
<given-names>P.</given-names>
</name>
</person-group> (<year>2023</year>). <article-title>Global distribution and morphology of small seamounts</article-title>. <source>Earth Space Sci.</source> <volume>10</volume> (<issue>4</issue>), <fpage>e2022EA002331</fpage>. <pub-id pub-id-type="doi">10.1029/2022ea002331</pub-id>
</citation>
</ref>
<ref id="B26">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Goode</surname>
<given-names>S. L.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Resilience of seamount benthic communities to trawling disturbance</article-title>. <source>Mar. Environ. Res.</source> <volume>161</volume>, <fpage>105086</fpage>. <pub-id pub-id-type="doi">10.1016/j.marenvres.2020.105086</pub-id>
<pub-id pub-id-type="pmid">32889447</pub-id>
</citation>
</ref>
<ref id="B27">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Goode</surname>
<given-names>S. L.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Stephenson</surname>
<given-names>F.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Fine-scale mapping of mega-epibenthic communities and their patch characteristics on two New Zealand seamounts</article-title>. <source>Front. Mar. Sci.</source> <volume>8</volume>, <fpage>765407</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2021.765407</pub-id>
</citation>
</ref>
<ref id="B28">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Greenwell</surname>
<given-names>B. M.</given-names>
</name>
</person-group> (<year>2017</year>). <article-title>pdp: an R package for constructing partial dependence plots</article-title>. <source>R. J.</source> <volume>9</volume> (<issue>1</issue>), <fpage>421</fpage>. <pub-id pub-id-type="doi">10.32614/rj-2017-016</pub-id>
</citation>
</ref>
<ref id="B29">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Gros</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Jansen</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Dunstan</surname>
<given-names>P. K.</given-names>
</name>
<name>
<surname>Welsford</surname>
<given-names>D. C.</given-names>
</name>
<name>
<surname>Hill</surname>
<given-names>N. A.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Vulnerable, but still poorly known, marine ecosystems: how to make distribution models more relevant and impactful for conservation and management of VMEs?</article-title> <source>Front. Mar. Sci.</source> <volume>9</volume>, <fpage>870145</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2022.870145</pub-id>
</citation>
</ref>
<ref id="B30">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Grosse</surname>
<given-names>P.</given-names>
</name>
<name>
<surname>Van Wyk De Vries</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Petrinovic</surname>
<given-names>I. A.</given-names>
</name>
<name>
<surname>Euillades</surname>
<given-names>P. A.</given-names>
</name>
<name>
<surname>Alvarado</surname>
<given-names>G. E.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>Morphometry and evolution of arc volcanoes</article-title>. <source>Geology</source> <volume>37</volume> (<issue>7</issue>), <fpage>651</fpage>&#x2013;<lpage>654</lpage>. <pub-id pub-id-type="doi">10.1130/g25734a.1</pub-id>
</citation>
</ref>
<ref id="B31">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Guinotte</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Davies</surname>
<given-names>A. J.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Predicted deep-sea coral habitat suitability for the U.S. West coast</article-title>. <source>PLoS ONE</source> <volume>9</volume> (<issue>4</issue>), <fpage>e93918</fpage>. <pub-id pub-id-type="doi">10.1371/journal.pone.0093918</pub-id>
<pub-id pub-id-type="pmid">24759613</pub-id>
</citation>
</ref>
<ref id="B32">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Guisan</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Tingley</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Baumgartner</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Naujokaitis-Lewis</surname>
<given-names>I.</given-names>
</name>
<name>
<surname>Sutcliffe</surname>
<given-names>P.</given-names>
</name>
<name>
<surname>Tulloch</surname>
<given-names>A.</given-names>
</name>
<etal/>
</person-group> (<year>2013</year>). <article-title>Predicting species distributions for conservation decisions</article-title>. <source>Ecol. Lett.</source> <volume>16</volume>, <fpage>1424</fpage>&#x2013;<lpage>1435</lpage>. <pub-id pub-id-type="doi">10.1111/ele.12189</pub-id>
<pub-id pub-id-type="pmid">24134332</pub-id>
</citation>
</ref>
<ref id="B33">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Hastie</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Tibshirani</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Friedman</surname>
<given-names>J. H.</given-names>
</name>
<name>
<surname>Friedman</surname>
<given-names>J. H.</given-names>
</name>
</person-group> (<year>2009</year>). <source>The elements of statistical learning: data mining, inference, and prediction</source>. <publisher-name>Springer</publisher-name>.</citation>
</ref>
<ref id="B34">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Heap</surname>
<given-names>A. D.</given-names>
</name>
<name>
<surname>Harris</surname>
<given-names>P. T.</given-names>
</name>
</person-group> (<year>2008</year>). <article-title>Geomorphology of the Australian margin and adjacent seafloor</article-title>. <source>Aust. J. Earth Sci.</source> <volume>55</volume> (<issue>4</issue>), <fpage>555</fpage>&#x2013;<lpage>585</lpage>. <pub-id pub-id-type="doi">10.1080/08120090801888669</pub-id>
</citation>
</ref>
<ref id="B35">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Henry</surname>
<given-names>L.-A.</given-names>
</name>
<name>
<surname>Roberts</surname>
<given-names>J. M.</given-names>
</name>
</person-group> (<year>2017</year>). <article-title>Global biodiversity in cold-water coral reef ecosystems</article-title>,&#x201d; in <source>Marine animal forests: the ecology of benthic biodiversity hotspots</source>, <fpage>235</fpage>&#x2013;<lpage>256</lpage>.</citation>
</ref>
<ref id="B36">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Hosmer</surname>
<given-names>D. W. J.</given-names>
</name>
<name>
<surname>Lemeshow</surname>
<given-names>S.</given-names>
</name>
<name>
<surname>Sturdivant</surname>
<given-names>R. X.</given-names>
</name>
</person-group> (<year>2013</year>). &#x201c;<article-title>Assessing the fit of the model</article-title>,&#x201d; in <source>Applied logistic regression</source>, <fpage>153</fpage>&#x2013;<lpage>225</lpage>.</citation>
</ref>
<ref id="B37">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Huang</surname>
<given-names>Z.</given-names>
</name>
<name>
<surname>Brooke</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Li</surname>
<given-names>J.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Performance of predictive models in marine benthic environments based on predictions of sponge distribution on the Australian continental shelf</article-title>. <source>Ecol. Inf.</source> <volume>6</volume> (<issue>3</issue>), <fpage>205</fpage>&#x2013;<lpage>216</lpage>. <pub-id pub-id-type="doi">10.1016/j.ecoinf.2011.01.001</pub-id>
</citation>
</ref>
<ref id="B38">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Koslow</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Gowlett-Holmes</surname>
<given-names>K.</given-names>
</name>
<name>
<surname>Lowry</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>O&#x27;Hara</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Poore</surname>
<given-names>G.</given-names>
</name>
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
</person-group> (<year>2001</year>). <article-title>Seamount benthic macrofauna off southern Tasmania: community structure and impacts of trawling</article-title>. <source>Mar. Ecol. Prog. Ser.</source> <volume>213</volume>, <fpage>111</fpage>&#x2013;<lpage>125</lpage>. <pub-id pub-id-type="doi">10.3354/meps213111</pub-id>
</citation>
</ref>
<ref id="B39">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lamarche</surname>
<given-names>G.</given-names>
</name>
<name>
<surname>Lurton</surname>
<given-names>X.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Recommendations for improved and coherent acquisition and processing of backscatter data from seafloor-mapping sonars</article-title>. <source>Mar. Geophys. Res.</source> <volume>39</volume> (<issue>1-2</issue>), <fpage>5</fpage>&#x2013;<lpage>22</lpage>. <pub-id pub-id-type="doi">10.1007/s11001-017-9315-6</pub-id>
</citation>
</ref>
<ref id="B40">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lamarche</surname>
<given-names>G.</given-names>
</name>
<name>
<surname>Lurton</surname>
<given-names>X.</given-names>
</name>
<name>
<surname>Verdier</surname>
<given-names>A.-L.</given-names>
</name>
<name>
<surname>Augustin</surname>
<given-names>J.-M.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Quantitative characterisation of seafloor substrate and bedforms using advanced processing of multibeam backscatter&#x2014;application to Cook Strait, New Zealand</article-title>. <source>Cont. Shelf Res.</source> <volume>31</volume> (<issue>2</issue>), <fpage>S93</fpage>&#x2013;<lpage>S109</lpage>. <pub-id pub-id-type="doi">10.1016/j.csr.2010.06.001</pub-id>
</citation>
</ref>
<ref id="B41">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Liaw</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Wiener</surname>
<given-names>M.</given-names>
</name>
</person-group> (<year>2001</year>). <article-title>Classification and regression by RandomForest</article-title>. <source>Forest</source> <volume>23</volume>.</citation>
</ref>
<ref id="B42">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lim</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Wheeler</surname>
<given-names>A. J.</given-names>
</name>
<name>
<surname>Conti</surname>
<given-names>L.</given-names>
</name>
</person-group> (<year>2020</year>). <article-title>Cold-water coral habitat mapping: trends and developments in acquisition and processing methods</article-title>. <source>Geosciences</source> <volume>11</volume> (<issue>1</issue>), <fpage>9</fpage>. <pub-id pub-id-type="doi">10.3390/geosciences11010009</pub-id>
</citation>
</ref>
<ref id="B43">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lucieer</surname>
<given-names>V.</given-names>
</name>
<name>
<surname>Lamarche</surname>
<given-names>G.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Unsupervised fuzzy classification and object-based image analysis of multibeam data to map deep water substrates, Cook Strait, New Zealand</article-title>. <source>Cont. Shelf Res.</source> <volume>31</volume> (<issue>11</issue>), <fpage>1236</fpage>&#x2013;<lpage>1247</lpage>. <pub-id pub-id-type="doi">10.1016/j.csr.2011.04.016</pub-id>
</citation>
</ref>
<ref id="B44">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lucieer</surname>
<given-names>V.</given-names>
</name>
<name>
<surname>Barrett</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>Butler</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Flukes</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Ierodiaconou</surname>
<given-names>D.</given-names>
</name>
<name>
<surname>Ingleton</surname>
<given-names>T.</given-names>
</name>
<etal/>
</person-group> (<year>2019</year>). <article-title>A seafloor habitat map for the Australian continental shelf</article-title>. <source>Sci. Data</source> <volume>6</volume> (<issue>1</issue>), <fpage>120</fpage>. <pub-id pub-id-type="doi">10.1038/s41597-019-0126-2</pub-id>
<pub-id pub-id-type="pmid">31296871</pub-id>
</citation>
</ref>
<ref id="B45">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Lurton</surname>
<given-names>X.</given-names>
</name>
<name>
<surname>Lamarche</surname>
<given-names>G.</given-names>
</name>
</person-group> (<year>2015</year>). <article-title>Backscatter measurements by seafloor-mapping sonars</article-title>. <source>Guidel. Recomm</source>. <comment>Available online at: <ext-link ext-link-type="uri" xlink:href="http://geohab.org/publications/">http://geohab.org/publications/</ext-link>.</comment>
</citation>
</ref>
<ref id="B46">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Maguire</surname>
<given-names>K.</given-names>
</name>
<name>
<surname>O&#x27;Neill</surname>
<given-names>H.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>White</surname>
<given-names>W.</given-names>
</name>
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
</person-group> (<year>2023</year>). <article-title>Seamount coral reefs are egg case nurseries for deep&#x2010;sea skates</article-title>. <source>J. Fish Biol.</source> <volume>102</volume> (<issue>6</issue>), <fpage>1455</fpage>&#x2013;<lpage>1469</lpage>. <pub-id pub-id-type="doi">10.1111/jfb.15376</pub-id>
<pub-id pub-id-type="pmid">36960821</pub-id>
</citation>
</ref>
<ref id="B47">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mayer</surname>
<given-names>L.</given-names>
</name>
<name>
<surname>Jakobsson</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Allen</surname>
<given-names>G.</given-names>
</name>
<name>
<surname>Dorschel</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Falconer</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Ferrini</surname>
<given-names>V.</given-names>
</name>
<etal/>
</person-group> (<year>2018</year>). <article-title>The nippon foundation&#x2014;GEBCO seabed 2030 project: the quest to see the world&#x2019;s oceans completely mapped by 2030</article-title>. <source>Geosciences</source> <volume>8</volume> (<issue>2</issue>), <fpage>63</fpage>. <pub-id pub-id-type="doi">10.3390/geosciences8020063</pub-id>
</citation>
</ref>
<ref id="B48">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Misiuk</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Brown</surname>
<given-names>C. J.</given-names>
</name>
</person-group> (<year>2022</year>). <article-title>Multiple imputation of multibeam angular response data for high resolution full coverage seabed mapping</article-title>. <source>Mar. Geophys. Res.</source> <volume>43</volume> (<issue>1</issue>), <fpage>7</fpage>. <pub-id pub-id-type="doi">10.1007/s11001-022-09471-3</pub-id>
</citation>
</ref>
<ref id="B49">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Mitchell</surname>
<given-names>G. A.</given-names>
</name>
<name>
<surname>Orange</surname>
<given-names>D. L.</given-names>
</name>
<name>
<surname>Gharib</surname>
<given-names>J. J.</given-names>
</name>
<name>
<surname>Kennedy</surname>
<given-names>P.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Improved detection and mapping of deepwater hydrocarbon seeps: optimizing multibeam echosounder seafloor backscatter acquisition and processing techniques</article-title>. <source>Mar. Geophys. Res.</source> <volume>39</volume>, <fpage>323</fpage>&#x2013;<lpage>347</lpage>. <pub-id pub-id-type="doi">10.1007/s11001-018-9345-8</pub-id>
</citation>
</ref>
<ref id="B50">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Pitcher</surname>
<given-names>T. J.</given-names>
</name>
<name>
<surname>Morato</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Hart</surname>
<given-names>P. J. B.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Hagaan</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>Santos</surname>
<given-names>R. S.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Seamounts: ecology, fisheries and conservation</article-title>. <pub-id pub-id-type="doi">10.1002/9780470691953</pub-id>
</citation>
</ref>
<ref id="B51">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Pittman</surname>
<given-names>S. J.</given-names>
</name>
<name>
<surname>Brown</surname>
<given-names>K. A.</given-names>
</name>
</person-group> (<year>2011</year>). <article-title>Multi-scale approach for predicting fish species distributions across coral reef seascapes</article-title>. <source>PLoS ONE</source> <volume>6</volume> (<issue>5</issue>), <fpage>e20583</fpage>. <pub-id pub-id-type="doi">10.1371/journal.pone.0020583</pub-id>
<pub-id pub-id-type="pmid">21637787</pub-id>
</citation>
</ref>
<ref id="B52">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Przeslawski</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Foster</surname>
<given-names>S.</given-names>
</name>
<name>
<surname>Monk</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Barrett</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>Bouchet</surname>
<given-names>P.</given-names>
</name>
<name>
<surname>Carroll</surname>
<given-names>A.</given-names>
</name>
<etal/>
</person-group> (<year>2019</year>). <article-title>A suite of field manuals for marine sampling to monitor Australian waters</article-title>. <source>Front. Mar. Sci.</source> <volume>6</volume>, <fpage>177</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2019.00177</pub-id>
</citation>
</ref>
<ref id="B53">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ramirez-Llodra</surname>
<given-names>E.</given-names>
</name>
<name>
<surname>Tyler</surname>
<given-names>P. A.</given-names>
</name>
<name>
<surname>Baker</surname>
<given-names>M. C.</given-names>
</name>
<name>
<surname>Bergstad</surname>
<given-names>O. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Escobar</surname>
<given-names>E.</given-names>
</name>
<etal/>
</person-group> (<year>2011</year>). <article-title>Man and the last great wilderness: human impact on the deep sea</article-title>. <source>PLoS ONE</source> <volume>6</volume> (<issue>8</issue>), <fpage>e22588</fpage>. <pub-id pub-id-type="doi">10.1371/journal.pone.0022588</pub-id>
<pub-id pub-id-type="pmid">21829635</pub-id>
</citation>
</ref>
<ref id="B54">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ramiro-S&#xe1;nchez</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Gonz&#xe1;lez-Irusta</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Henry</surname>
<given-names>L.-A.</given-names>
</name>
<name>
<surname>Cleland</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Yeo</surname>
<given-names>I.</given-names>
</name>
<name>
<surname>Xavier</surname>
<given-names>J. R.</given-names>
</name>
<etal/>
</person-group> (<year>2019</year>). <article-title>Characterization and mapping of a deep-sea sponge ground on the tropic seamount (northeast tropical atlantic): implications for spatial management in the high seas</article-title>. <source>Front. Mar. Sci.</source> <volume>6</volume>, <fpage>278</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2019.00278</pub-id>
</citation>
</ref>
<ref id="B55">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rengstorf</surname>
<given-names>A. M.</given-names>
</name>
<name>
<surname>Grehan</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Yesson</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Brown</surname>
<given-names>C.</given-names>
</name>
</person-group> (<year>2012</year>). <article-title>Towards high-resolution habitat suitability modeling of vulnerable marine ecosystems in the deep-sea: resolving terrain attribute dependencies</article-title>. <source>Mar. Geod.</source> <volume>35</volume> (<issue>4</issue>), <fpage>343</fpage>&#x2013;<lpage>361</lpage>. <pub-id pub-id-type="doi">10.1080/01490419.2012.699020</pub-id>
</citation>
</ref>
<ref id="B56">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Roberts</surname>
<given-names>J. M.</given-names>
</name>
</person-group> (<year>2009</year>). <source>Cold-water corals: the biology and geology of deep-sea coral habitats</source>. <publisher-name>Cambridge University Press</publisher-name>.</citation>
</ref>
<ref id="B57">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Roberts</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Cairns</surname>
<given-names>S. D.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Cold-water corals in a changing ocean</article-title>. <source>Curr. Opin. Environ. Sustain.</source> <volume>7</volume>, <fpage>118</fpage>&#x2013;<lpage>126</lpage>. <pub-id pub-id-type="doi">10.1016/j.cosust.2014.01.004</pub-id>
</citation>
</ref>
<ref id="B58">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Roberts</surname>
<given-names>D. R.</given-names>
</name>
<name>
<surname>Bahn</surname>
<given-names>V.</given-names>
</name>
<name>
<surname>Ciuti</surname>
<given-names>S.</given-names>
</name>
<name>
<surname>Boyce</surname>
<given-names>M. S.</given-names>
</name>
<name>
<surname>Elith</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Guillera&#x2010;Arroita</surname>
<given-names>G.</given-names>
</name>
<etal/>
</person-group> (<year>2017</year>). <article-title>Cross&#x2010;validation strategies for data with temporal, spatial, hierarchical, or phylogenetic structure</article-title>. <source>Ecography</source> <volume>40</volume> (<issue>8</issue>), <fpage>913</fpage>&#x2013;<lpage>929</lpage>. <pub-id pub-id-type="doi">10.1111/ecog.02881</pub-id>
</citation>
</ref>
<ref id="B59">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Rogers</surname>
<given-names>A. D.</given-names>
</name>
</person-group> (<year>2018</year>). &#x201c;<article-title>Chapter four - the biology of seamounts: 25 Years on</article-title>,&#x201d; in <source>Advances in marine biology</source>. Editor <person-group person-group-type="editor">
<name>
<surname>Sheppard</surname>
<given-names>C.</given-names>
</name>
</person-group> (<publisher-name>Academic Press</publisher-name>), <fpage>137</fpage>&#x2013;<lpage>224</lpage>.</citation>
</ref>
<ref id="B60">
<citation citation-type="book">
<person-group person-group-type="author">
<name>
<surname>Rogers</surname>
<given-names>A. D.</given-names>
</name>
</person-group> (<year>2019</year>). &#x201c;<article-title>Chapter 23 - threats to seamount ecosystems and their management</article-title>,&#x201d; in <source>World seas: an environmental evaluation</source>. Editor <person-group person-group-type="editor">
<name>
<surname>Sheppard</surname>
<given-names>C.</given-names>
</name>
</person-group> <edition>Second Edition</edition> (<publisher-name>Academic Press</publisher-name>), <fpage>427</fpage>&#x2013;<lpage>451</lpage>.</citation>
</ref>
<ref id="B61">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Georgian</surname>
<given-names>S. E.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Pallentin</surname>
<given-names>A.</given-names>
</name>
<etal/>
</person-group> (<year>2017</year>). <article-title>High-resolution habitat suitability models for the conservation and management of vulnerable marine ecosystems on the Louisville seamount chain, south pacific ocean</article-title>. <source>Front. Mar. Sci.</source> <volume>4</volume>, <fpage>335</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2017.00335</pub-id>
</citation>
</ref>
<ref id="B62">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Ryabinin</surname>
<given-names>V.</given-names>
</name>
<name>
<surname>Barbi&#xe8;re</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Haugan</surname>
<given-names>P.</given-names>
</name>
<name>
<surname>Kullenberg</surname>
<given-names>G.</given-names>
</name>
<name>
<surname>Smith</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>McLean</surname>
<given-names>C.</given-names>
</name>
<etal/>
</person-group> (<year>2019</year>). <article-title>The UN decade of Ocean science for sustainable development</article-title>. <source>Front. Mar. Sci.</source> <volume>6</volume>, <fpage>470</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2019.00470</pub-id>
</citation>
</ref>
<ref id="B63">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Schimel</surname>
<given-names>A. C. G.</given-names>
</name>
<name>
<surname>Beaudoin</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Parnum</surname>
<given-names>I. M.</given-names>
</name>
<name>
<surname>Le Bas</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Schmidt</surname>
<given-names>V.</given-names>
</name>
<name>
<surname>Keith</surname>
<given-names>G.</given-names>
</name>
<etal/>
</person-group> (<year>2018</year>). <article-title>Multibeam sonar backscatter data processing</article-title>. <source>Mar. Geophys. Res.</source> <volume>39</volume> (<issue>1-2</issue>), <fpage>121</fpage>&#x2013;<lpage>137</lpage>. <pub-id pub-id-type="doi">10.1007/s11001-018-9341-z</pub-id>
</citation>
</ref>
<ref id="B64">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Schlacher</surname>
<given-names>T. A.</given-names>
</name>
<name>
<surname>Baco</surname>
<given-names>A. R.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>O&#x27;Hara</surname>
<given-names>T. D.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Kelley</surname>
<given-names>C.</given-names>
</name>
<etal/>
</person-group> (<year>2014</year>). <article-title>Seamount benthos in a cobalt&#x2010;rich crust region of the central Pacific: conservation challenges for future seabed mining</article-title>. <source>Divers. Distributions</source> <volume>20</volume> (<issue>5</issue>), <fpage>491</fpage>&#x2013;<lpage>502</lpage>. <pub-id pub-id-type="doi">10.1111/ddi.12142</pub-id>
</citation>
</ref>
<ref id="B65">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Staudigel</surname>
<given-names>H.</given-names>
</name>
<name>
<surname>Clague</surname>
<given-names>D. A.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>The geological history of deep-sea volcanoes: biosphere, hydrosphere, and lithosphere interactions</article-title>. <source>Oceanography</source> <volume>23</volume> (<issue>1</issue>), <fpage>58</fpage>&#x2013;<lpage>71</lpage>. <pub-id pub-id-type="doi">10.5670/oceanog.2010.62</pub-id>
</citation>
</ref>
<ref id="B66">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Stephenson</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Pitcher</surname>
<given-names>C. R.</given-names>
</name>
<name>
<surname>Pinkerton</surname>
<given-names>M. H.</given-names>
</name>
<name>
<surname>Petersen</surname>
<given-names>G.</given-names>
</name>
<etal/>
</person-group> (<year>2021</year>). <article-title>Presence-only habitat suitability models for vulnerable marine ecosystem indicator taxa in the South Pacific have reached their predictive limit</article-title>. <source>ICES J. Mar. Sci.</source> <volume>78</volume> (<issue>8</issue>), <fpage>2830</fpage>&#x2013;<lpage>2843</lpage>. <pub-id pub-id-type="doi">10.1093/icesjms/fsab162</pub-id>
</citation>
</ref>
<ref id="B67">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Stephenson</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Anderson</surname>
<given-names>O. F.</given-names>
</name>
<name>
<surname>Ellis</surname>
<given-names>J. I.</given-names>
</name>
<name>
<surname>Geange</surname>
<given-names>S. W.</given-names>
</name>
<name>
<surname>Brough</surname>
<given-names>T.</given-names>
</name>
<etal/>
</person-group> (<year>2023</year>). <article-title>Implications for the conservation of deep-water corals in the face of multiple stressors: a case study from the New Zealand region</article-title>. <source>J. Environ. Manag.</source> <volume>346</volume>, <fpage>118938</fpage>. <pub-id pub-id-type="doi">10.1016/j.jenvman.2023.118938</pub-id>
<pub-id pub-id-type="pmid">37738731</pub-id>
</citation>
</ref>
<ref id="B68">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Strobl</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Malley</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Tutz</surname>
<given-names>G.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>An introduction to recursive partitioning: rationale, application, and characteristics of classification and regression trees, bagging, and random forests</article-title>. <source>Psychol. Methods</source> <volume>14</volume> (<issue>4</issue>), <fpage>323</fpage>&#x2013;<lpage>348</lpage>. <pub-id pub-id-type="doi">10.1037/a0016973</pub-id>
<pub-id pub-id-type="pmid">19968396</pub-id>
</citation>
</ref>
<ref id="B69">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Telford</surname>
<given-names>R. J.</given-names>
</name>
<name>
<surname>Birks</surname>
<given-names>H. J. B.</given-names>
</name>
</person-group> (<year>2009</year>). <article-title>Evaluation of transfer functions in spatially structured environments</article-title>. <source>Quat. Sci. Rev.</source> <volume>28</volume> (<issue>13</issue>), <fpage>1309</fpage>&#x2013;<lpage>1316</lpage>. <pub-id pub-id-type="doi">10.1016/j.quascirev.2008.12.020</pub-id>
</citation>
</ref>
<ref id="B70">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Thurber</surname>
<given-names>A. R.</given-names>
</name>
<name>
<surname>Sweetman</surname>
<given-names>A. K.</given-names>
</name>
<name>
<surname>Narayanaswamy</surname>
<given-names>B. E.</given-names>
</name>
<name>
<surname>Jones</surname>
<given-names>D. O. B.</given-names>
</name>
<name>
<surname>Ingels</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Hansman</surname>
<given-names>R. L.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Ecosystem function and services provided by the deep sea</article-title>. <source>Biogeosciences</source> <volume>11</volume> (<issue>14</issue>), <fpage>3941</fpage>&#x2013;<lpage>3963</lpage>. <pub-id pub-id-type="doi">10.5194/bg-11-3941-2014</pub-id>
</citation>
</ref>
<ref id="B71">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Tittensor</surname>
<given-names>D. P.</given-names>
</name>
<name>
<surname>Baco</surname>
<given-names>A. R.</given-names>
</name>
<name>
<surname>Hall-Spencer</surname>
<given-names>J. M.</given-names>
</name>
<name>
<surname>Orr</surname>
<given-names>J. C.</given-names>
</name>
<name>
<surname>Rogers</surname>
<given-names>A. D.</given-names>
</name>
</person-group> (<year>2010</year>). <article-title>Seamounts as refugia from ocean acidification for cold-water stony corals</article-title>. <source>Mar. Ecol.</source> <volume>31</volume> (<issue>s1</issue>), <fpage>212</fpage>&#x2013;<lpage>225</lpage>. <pub-id pub-id-type="doi">10.1111/j.1439-0485.2010.00393.x</pub-id>
</citation>
</ref>
<ref id="B72">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Tobler</surname>
<given-names>W. R.</given-names>
</name>
</person-group> (<year>1970</year>). <article-title>A computer movie simulating urban growth in the Detroit region</article-title>. <source>Econ. Geogr.</source> <volume>46</volume> (<issue>Suppl. 1</issue>), <fpage>234</fpage>&#x2013;<lpage>240</lpage>. <pub-id pub-id-type="doi">10.2307/143141</pub-id>
</citation>
</ref>
<ref id="B73">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Townsend</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>Baldry</surname>
<given-names>K.</given-names>
</name>
<name>
<surname>Crossman</surname>
<given-names>D.</given-names>
</name>
<name>
<surname>Cullen</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Doubell</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Ingleton</surname>
<given-names>T.</given-names>
</name>
<etal/>
</person-group> (<year>2023</year>). <article-title>Ausseabed: a national program of collaboration to maximise Australia&#x2019;s seabed mapping efforts</article-title>. <source>Coast. Eng. Proc.</source> <volume>37</volume>, <fpage>174</fpage>. <pub-id pub-id-type="doi">10.9753/icce.v37.management.174</pub-id>
</citation>
</ref>
<ref id="B74">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Valavi</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Elith</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Lahoz&#x2010;Monfort</surname>
<given-names>J. J.</given-names>
</name>
<name>
<surname>Guillera&#x2010;Arroita</surname>
<given-names>G.</given-names>
</name>
</person-group> (<year>2019</year>). <article-title>blockCV: an r package for generating spatially or environmentally separated folds for <italic>k</italic>&#x2010;fold cross&#x2010;validation of species distribution models</article-title>. <source>Methods Ecol. Evol.</source> <volume>10</volume> (<issue>2</issue>), <fpage>225</fpage>&#x2013;<lpage>232</lpage>. <pub-id pub-id-type="doi">10.1111/2041-210x.13107</pub-id>
</citation>
</ref>
<ref id="B75">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Valavi</surname>
<given-names>R.</given-names>
</name>
<name>
<surname>Elith</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Lahoz&#x2010;Monfort</surname>
<given-names>J. J.</given-names>
</name>
<name>
<surname>Guillera&#x2010;Arroita</surname>
<given-names>G.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>Modelling species presence&#x2010;only data with random forests</article-title>. <source>Ecography</source> <volume>44</volume> (<issue>12</issue>), <fpage>1731</fpage>&#x2013;<lpage>1742</lpage>. <pub-id pub-id-type="doi">10.1111/ecog.05615</pub-id>
</citation>
</ref>
<ref id="B76">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Vierod</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Guinotte</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Davies</surname>
<given-names>A.</given-names>
</name>
</person-group> (<year>2014</year>). <article-title>Predicting the distribution of vulnerable marine ecosystems in the deep sea using presence-background models</article-title>. <source>Deep Sea Res. Part II Top. Stud. Oceanogr.</source> <volume>99</volume>, <fpage>6</fpage>&#x2013;<lpage>18</lpage>. <pub-id pub-id-type="doi">10.1016/j.dsr2.2013.06.010</pub-id>
</citation>
</ref>
<ref id="B77">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Walbridge</surname>
<given-names>S.</given-names>
</name>
<name>
<surname>Slocum</surname>
<given-names>N.</given-names>
</name>
<name>
<surname>Pobuda</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Wright</surname>
<given-names>D.</given-names>
</name>
</person-group> (<year>2018</year>). <article-title>Unified geomorphological analysis workflows with benthic terrain modeler</article-title>. <source>Geosciences</source> <volume>8</volume> (<issue>3</issue>), <fpage>94</fpage>. <pub-id pub-id-type="doi">10.3390/geosciences8030094</pub-id>
</citation>
</ref>
<ref id="B78">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Wessel</surname>
<given-names>P.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Seamount characteristics</article-title>. <source>Seamounts Ecol. Fish. and conservation</source>, <fpage>1</fpage>&#x2013;<lpage>25</lpage>. <pub-id pub-id-type="doi">10.1002/9780470691953.ch1</pub-id>
</citation>
</ref>
<ref id="B79">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>White</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Bashmachnikov</surname>
<given-names>I.</given-names>
</name>
<name>
<surname>Ar&#xed;stegui</surname>
<given-names>J.</given-names>
</name>
<name>
<surname>Martins</surname>
<given-names>A.</given-names>
</name>
</person-group> (<year>2007</year>). <article-title>Physical processes and seamount productivity</article-title>. <source>Seamounts Ecol. Fish. and conservation</source>, <fpage>62</fpage>&#x2013;<lpage>84</lpage>. <pub-id pub-id-type="doi">10.1002/9780470691953.ch4</pub-id>
</citation>
</ref>
<ref id="B80">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Schlacher</surname>
<given-names>T. A.</given-names>
</name>
<name>
<surname>Rowden</surname>
<given-names>A. A.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Clark</surname>
<given-names>M. R.</given-names>
</name>
<name>
<surname>Bowden</surname>
<given-names>D. A.</given-names>
</name>
<etal/>
</person-group> (<year>2010</year>). <article-title>Seamount megabenthic assemblages fail to recover from trawling impacts</article-title>. <source>Mar. Ecol.</source> <volume>31</volume>, <fpage>183</fpage>&#x2013;<lpage>199</lpage>. <pub-id pub-id-type="doi">10.1111/j.1439-0485.2010.00385.x</pub-id>
</citation>
</ref>
<ref id="B81">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Green</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Maguire</surname>
<given-names>K.</given-names>
</name>
<name>
<surname>Untiedt</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Mortimer</surname>
<given-names>N.</given-names>
</name>
<etal/>
</person-group> (<year>2020a</year>). <article-title>True size matters for conservation: a robust method to determine the size of deep-sea coral reefs shows they are typically small on seamounts in the southwest pacific ocean</article-title>. <source>Front. Mar. Sci.</source> <volume>7</volume>, <fpage>187</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2020.00187</pub-id>
</citation>
</ref>
<ref id="B82">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Maguire</surname>
<given-names>K.</given-names>
</name>
<name>
<surname>Green</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Untiedt</surname>
<given-names>C.</given-names>
</name>
<name>
<surname>Alderslade</surname>
<given-names>P.</given-names>
</name>
<etal/>
</person-group> (<year>2020b</year>). <article-title>The fate of deep-sea coral reefs on seamounts in a fishery-seascape: what are the impacts, what remains, and what is protected?</article-title> <source>Front. Mar. Sci.</source> <volume>7</volume>, <fpage>567002</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2020.567002</pub-id>
</citation>
</ref>
<ref id="B83">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Williams</surname>
<given-names>A.</given-names>
</name>
<name>
<surname>Osterhage</surname>
<given-names>D.</given-names>
</name>
<name>
<surname>Althaus</surname>
<given-names>F.</given-names>
</name>
<name>
<surname>Ryan</surname>
<given-names>T.</given-names>
</name>
<name>
<surname>Green</surname>
<given-names>M.</given-names>
</name>
<name>
<surname>Pogonoski</surname>
<given-names>J.</given-names>
</name>
</person-group> (<year>2021</year>). <article-title>A very large spawning aggregation of a deep-sea eel: magnitude and status</article-title>. <source>J. Mar. Sci. Eng.</source> <volume>9</volume> (<issue>7</issue>), <fpage>723</fpage>. <pub-id pub-id-type="doi">10.3390/jmse9070723</pub-id>
</citation>
</ref>
<ref id="B84">
<citation citation-type="journal">
<person-group person-group-type="author">
<name>
<surname>Winship</surname>
<given-names>A. J.</given-names>
</name>
<name>
<surname>Thorson</surname>
<given-names>J. T.</given-names>
</name>
<name>
<surname>Clarke</surname>
<given-names>M. E.</given-names>
</name>
<name>
<surname>Coleman</surname>
<given-names>H. M.</given-names>
</name>
<name>
<surname>Costa</surname>
<given-names>B.</given-names>
</name>
<name>
<surname>Georgian</surname>
<given-names>S. E.</given-names>
</name>
<etal/>
</person-group> (<year>2020</year>). <article-title>Good practices for species distribution modeling of deep-sea corals and sponges for resource management: data collection, analysis, validation, and communication</article-title>. <source>Front. Mar. Sci.</source> <volume>7</volume>, <fpage>303</fpage>. <pub-id pub-id-type="doi">10.3389/fmars.2020.00303</pub-id>
</citation>
</ref>
</ref-list>
</back>
</article>