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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">118</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:71cc5dc6-a767-5334-951f-ef6ae8936459</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Plant Ecology and Evolution</journal-title>
        <abbrev-journal-title xml:lang="en">plecevo</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">2032-3913</issn>
      <issn pub-type="epub">2032-3921</issn>
      <publisher>
        <publisher-name>Meise Botanic Garden and Royal Botanical Society of Belgium</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.5091/plecevo.122470</article-id>
      <article-id pub-id-type="publisher-id">122470</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Angiospermae</subject>
          <subject>Fabaceae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
          <subject>Biogeography</subject>
          <subject>Data analysis &amp; Modelling</subject>
          <subject>Ecology</subject>
          <subject>Global Change</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Africa</subject>
          <subject>West Africa</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Past, present, and future potential distributions of the African multipurpose tree <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Fabaceae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Dassou</surname>
            <given-names>Gbèwonmèdéa Hospice</given-names>
          </name>
          <email xlink:type="simple">daspice2@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-4651-1802</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <xref ref-type="aff" rid="A2">2</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/project-administration/">Project administration</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Agoundé</surname>
            <given-names>Gafarou</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/formal-analysis/">Formal analysis</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/software/">Software</role>
          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Akouété</surname>
            <given-names>Pathmos</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
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          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Favi</surname>
            <given-names>Gnimansou Abraham</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-6243-9236</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
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          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kpétikou</surname>
            <given-names>Ghyslain Chabi</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
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          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Salako</surname>
            <given-names>Kolawolé Valère</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-7817-3687</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
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          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Ouachinou</surname>
            <given-names>Jéronime Marie-Ange Sènami</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-9940-5061</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Makponsè</surname>
            <given-names>Judicael</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/data-curation/">Data curation</role>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Kouyaté</surname>
            <given-names>Amadou Malé</given-names>
          </name>
          <xref ref-type="aff" rid="A4">4</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Sari</surname>
            <given-names>İdris</given-names>
          </name>
          <xref ref-type="aff" rid="A5">5</xref>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Glèlè Kakaï</surname>
            <given-names>Romain Lucas</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Yédomonhan</surname>
            <given-names>Hounnankpon</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Adomou</surname>
            <given-names>Aristide Cossi</given-names>
          </name>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-review-editing/">Writing - review and editing</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</role>
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      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Herbier National du Bénin, Laboratoire des Sciences du Végétal et Pharmacopée, École Doctorale des Sciences de la Vie et de la Terre, Université d’Abomey-Calavi, Cotonou, Benin</addr-line>
        <institution>National Herbarium of Benin</institution>
        <addr-line content-type="city">Abomey-Calavi</addr-line>
        <country>Benin</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Jardins Botanique et Zoologique Edouard Adjanohoun, Université d’Abomey-Calavi, Cotonou, Benin</addr-line>
        <institution>Botanical and Zoological Garden Edouard Adjanohoun</institution>
        <addr-line content-type="city">Abomey-Calavi</addr-line>
        <country>Benin</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Laboratoire de Biomathématiques et d’Estimations Forestières, École Doctorale des Sciences Agronomiques et de l’Eau, Université d’Abomey-Calavi, Cotonou, Benin</addr-line>
        <institution>Université d’Abomey-Calavi</institution>
        <addr-line content-type="city">Abomey-Calavi</addr-line>
        <country>Benin</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Département des Sciences Biologiques Appliquées, Centre Régional de la Recherche Agronomique, Institut d’Économie Rurale, Sikasso, Mali</addr-line>
        <institution>Centre Régional de la Recherche Agronomique, Institut d’Économie Rurale</institution>
        <addr-line content-type="city">Sikasso</addr-line>
        <country>Mali</country>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">Department of Biology, Faculty of Sciences and Letters, Erzincan Binali Yildirim University, Erzincan, Turkey</addr-line>
        <institution>Erzincan Binali Yildirim University</institution>
        <addr-line content-type="city">Erzincan</addr-line>
        <country>Turkiye</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Gbèwonmèdéa Hospice Dassou (<email xlink:type="simple">daspice2@gmail.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Brecht Verstraete</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2024</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>25</day>
        <month>09</month>
        <year>2024</year>
      </pub-date>
      <volume>157</volume>
      <issue>3</issue>
      <fpage>343</fpage>
      <lpage>357</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/452A92CD-41B9-51D8-B78A-C3ACCBACE075">452A92CD-41B9-51D8-B78A-C3ACCBACE075</uri>
      <history>
        <date date-type="received">
          <day>07</day>
          <month>03</month>
          <year>2024</year>
        </date>
        <date date-type="accepted">
          <day>29</day>
          <month>07</month>
          <year>2024</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Gbèwonmèdéa Hospice Dassou, Gafarou Agoundé, Pathmos Akouété, Gnimansou Abraham Favi, Ghyslain Chabi Kpétikou, Kolawolé Valère Salako, Jéronime Marie-Ange Sènami Ouachinou, Judicael Makponsè, Amadou Malé Kouyaté, İdris Sari, Romain Lucas Glèlè Kakaï, Hounnankpon Yédomonhan, Aristide Cossi Adomou</copyright-statement>
        <license license-type="creative-commons-attribution" xlink:href="http://creativecommons.org/licenses/by/4.0/" xlink:type="simple">
          <license-p>This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.</license-p>
        </license>
      </permissions>
      <abstract>
        <label>Abstract</label>
        <p><bold>Background and aims</bold> – Climate change induces increasing temperatures and drought, with possible profound shifts in species’ presence and distribution. Ecological niche models are widely used to assess plant species responses to climate change. However, such data are scarce for West Africa, particularly for vulnerable multipurpose species. This study focuses on modelling the ecological niche and the conservation status of the multipurpose tree <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> to improve insights into its habitat suitability in West Africa under past, present, and future climatic conditions. This will provide an essential basis for setting up global management plans through efficient conservation and ecological restoration policies.</p>
        <p><bold>Material and methods</bold> – The potential distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> under past, current, and future climate scenarios were assessed using four algorithms including generalized additive models (<abbrev xlink:title="generalized additive models" id="ABBRID0ESAAC">GAM</abbrev>), generalized linear models (<abbrev xlink:title="generalized linear models" id="ABBRID0EWAAC">GLM</abbrev>), random forest (<abbrev xlink:title="random forest" id="ABBRID0E1AAC">RF</abbrev>), and Maximum Entropy (<abbrev xlink:title="Maximum Entropy" id="ABBRID0E5AAC">MaxEnt</abbrev>). We also assessed the shift direction of suitable habitats and the conservation status of the species based on IUCN criteria. Overall, 220 occurrences were combined with a set of five bioclimatic variables to run the models.</p>
        <p><bold>Key results</bold> – Models performed well with good values of <abbrev xlink:title="operating characteristic curve" id="ABBRID0EGBAC">AUC</abbrev> (0.92) and <abbrev xlink:title="true skills statistic" id="ABBRID0EKBAC">TSS</abbrev> (0.73). Isothermality (41.10%) and Precipitation of Wettest Month (21.50%) contributed most to the distribution of the species. The distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> was relatively constant from the past to the present but could decrease in the next decades. In the future, only 17.70% and 13.98% of the areas were predicted to be suitable under respectively ssp245 and ssp585. In protected areas, the suitable areas under ssp245 were estimated at 21.01% with a decrease of 2.50% and 14.60% with a decrease of 8.61% under ssp585 by 2050. The direction of the distribution shifted to the south-east under future climate scenarios. The conservation status assessment of the species is Least Concern (<abbrev xlink:title="Least Concern" id="ABBRID0EZBAC">LC</abbrev>).</p>
        <p><bold>Conclusion</bold> – This study improves our understanding of the past, present-day, and future distribution of the species and provides support to better manage the conservation of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in West Africa.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>centroid</kwd>
        <kwd>climate</kwd>
        <kwd>conservation</kwd>
        <kwd>
          <italic>
            <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name>
          </italic>
        </kwd>
        <kwd>ecological niche modelling</kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>International Foundation for Science (IFS)</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="SECID0EDDAC">
      <title>Introduction</title>
      <p>The genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part></tp:taxon-name></italic> Juss. (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Fabaceae</tp:taxon-name-part></tp:taxon-name>) consists of three species found from West Tropical Africa to Sudan and it is introduced in Trinidad and Tobago (<xref ref-type="bibr" rid="B84">WFO 2024</xref>). Among them, the Tallow tree (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> J.F.Gmel.) is probably one of the most known <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part></tp:taxon-name></italic> species. It has a great socio-economic value in West Africa and has the potential to contribute to food security. The species ranks among the highly valued non-timber forest products for its nutritional and medicinal value, and magical and therapeutic uses throughout West Africa (<xref ref-type="bibr" rid="B18">Dassou et al. 2023</xref>). The nutritional and medicinal value of the fruit pulp and seed oil are exceptional. For instance, the pulp is rich in vitamin C, with more than 1000 mg / 100 g (<xref ref-type="bibr" rid="B21">Diop et al. 2010</xref>), which is about 29, 12, 7, and 5 times higher than in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Citrus">Citrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="hybrid-sign">×</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="sinensis">sinensis</tp:taxon-name-part></tp:taxon-name></italic> (L.) Osbeck, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Citrus">Citrus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="hybrid-sign">×</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="limon">limon</tp:taxon-name-part></tp:taxon-name></italic> (L.) Osbeck, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Psidium">Psidium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="guajava">guajava</tp:taxon-name-part></tp:taxon-name></italic> L., and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Adansonia">Adansonia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="digitata">digitata</tp:taxon-name-part></tp:taxon-name></italic> L. pulps respectively. It also contains vitamins B1 (thiamin), B2 (riboflavin), B3 (niacin), provitamine A (β-carotene), and essential amino acids (<xref ref-type="bibr" rid="B58">National Research Council 2008</xref>; <xref ref-type="bibr" rid="B21">Diop et al. 2010</xref>; <xref ref-type="bibr" rid="B59">Ndiaye et al. 2016</xref>). The seeds contain various minerals such as potassium (99.26 mg/g), magnesium (77.83 mg/g), calcium (71.11 mg/g), sodium (55.26 mg/g), iron (30.21 mg/g), manganese (7.89 mg/g), zinc (5.26 mg/g), and copper (4.99 mg/g) (<xref ref-type="bibr" rid="B74">Sowemimo et al. 2011</xref>). The leaves, bark, roots, pulps, and seeds are rich in therapeutic substances, including powerful anti-bacterial, anti-viral, hypoglycemic, and anti-cancer molecules (<xref ref-type="bibr" rid="B6">Akah et al. 2012</xref>; <xref ref-type="bibr" rid="B70">Sanni et al. 2015</xref>).</p>
      <p>Unfortunately, in addition to the threat of habitat degradation and its exploitation as timber, there is a growing regional trade of its fruits and seeds (<xref ref-type="bibr" rid="B60">Neuenschwander et al. 2011</xref>; <xref ref-type="bibr" rid="B42">Houénon et al. 2021</xref>). It has been suggested that the high commercial value of the fruits and seeds has stimulated its collection to a level which may threaten the regeneration of the species. Because of this persistent over-exploitation, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> has been reported as a threatened species in Benin (“Endangered”) (<xref ref-type="bibr" rid="B3">Adomou et al. 2009</xref>; <xref ref-type="bibr" rid="B60">Neuenschwander et al. 2011</xref>) as well as in West Africa (<xref ref-type="bibr" rid="B36">Hawthorne 1996</xref>). Moreover, the rate of natural regeneration from seeds, even after dispersal by mammals, is very low (<xref ref-type="bibr" rid="B22">El-Kamali 2011</xref>; <xref ref-type="bibr" rid="B41">Houénon et al. 2022</xref>). As such, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> risks, in the medium-term, to become very scarce or even disappear locally, although it is classified as “Least Concern” on the IUCN Red List (<xref ref-type="bibr" rid="B11">BGCI and IUCN 2019</xref>). While the cultivation of this precious fruit tree is urgent, it is equally important to engage in sustainable conservation strategies to preserve its natural populations. One of the key variables to consider both in cultivation and conservation projects is the species’ responses to climatic variation. Yet, it is still unclear how various climate changes influenced its distribution and how potential areas for its cultivation can change across its distribution range.</p>
      <p>Understanding the distribution of key taxa and their response to environmental change is fundamental to their effective management (<xref ref-type="bibr" rid="B1">Acevedo et al. 2012</xref>). In West Africa, patterns of distribution around the Dahomey Gap have been of particular interest to palynologists, archaeologists, and biogeographers as they can help to explain present-day distribution of the Guineo-Congolian closed semi-deciduous humid forests limited to tiny patches, most of them so-called sacred forests in Benin, Togo, and Ghana (<xref ref-type="bibr" rid="B79">Tossou 2002</xref>; <xref ref-type="bibr" rid="B2">Adomou 2005</xref>; <xref ref-type="bibr" rid="B69">Salzmann and Hoelzmann 2005</xref>). The establishment of the Dahomey Gap is favoured by the combined effect of the orientation of the coast in relation to sea currents, which locally modify the climate of the Gulf of Benin, climate change from the Quaternary (especially the Pleistocene and the Holocene), the harmattan, and human activities notably agriculture and forest exploitation (<xref ref-type="bibr" rid="B69">Salzmann and Hoelzmann 2005</xref>). Due to these factors, West Africa’s rainforests receded into two major blocks: the western Upper Guinean situated in today’s Ivory Coast/Liberia, and the eastern Lower Guinean/Congolese forest block in southern Cameroon, which are separated by the Dahomey Gap where the savannah reaches the coast (<xref ref-type="bibr" rid="B66">Poorter et al. 2004</xref>; <xref ref-type="bibr" rid="B69">Salzmann and Hoelzmann 2005</xref>; <xref ref-type="bibr" rid="B27">Giresse 2007</xref>). Consequently, in southern Benin and Togo, dense humid forest only exists in the form of islands or relics and the evergreen dense forest unit of dry land is completely absent. Indirectly, the current ﻿distribution pattern of forest species in the Dahomey Gap is not only determined by current climatic gradients but also by Quaternary climatic changes. Understanding the specific properties of these changes, which may have an impact on species or their habitats, is central to the sustainable management of species and their ecosystems (<xref ref-type="bibr" rid="B37">Heller and Zavaleta 2009</xref>). These climate changes constitute an environmental issue that deserves special attention in the planning of agricultural production, the diversification of agricultural production, and the conservation of species. Fluctuations in climatic variables such as precipitation and temperature are increasingly likely to affect biodiversity and the geographic distribution of favourable habitats for species (<xref ref-type="bibr" rid="B46">IPCC 2007</xref>). Information on the impact of climate change on biodiversity is also important to better reason the choices of areas where species can be conserved today and in the future. This is because climate change could generate a spatial dynamic in the geographical distribution of habitats favourable to species and thus make certain regions favourable today, while very unfavourable in the future and vice versa (<xref ref-type="bibr" rid="B34">Hannah et al. 2002</xref>). Overlooking this dimension could jeopardize efforts to conserve and cultivate the species.</p>
      <p>Species distribution modelling (<abbrev xlink:title="Species distribution modelling" id="ABBRID0EFKAC">SDM</abbrev>) is one of the commonly used tools for investigating potential geographical distributions of organisms. <abbrev xlink:title="Species distribution modelling" id="ABBRID0EJKAC">SDM</abbrev> can be defined as the prediction of species distribution across the landscape based on the relationship between species occurrence and environmental variables (<xref ref-type="bibr" rid="B32">Guisan et al. 2002</xref>). Such distribution models are becoming important tools for survey design and the most popular is the Maximum Entropy (<abbrev xlink:title="Maximum Entropy" id="ABBRID0ERKAC">MaxEnt</abbrev>). Modelling the ecological niche and conservation potential of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in protected areas under historical, contemporary, and projected West African climates is the main goal of this research. Specifically, this study aims to: (i) identify the environmental variables that significantly influence the distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in West Africa; (ii) determine the potential habitats favourable to the geographical distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> under past, present, and future climatic conditions; (iii) assess the shift direction in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution in West Africa under past, current, and future climates; and (iv) assess the changes of habitat suitability in the current and future distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in existent protected areas. We further discuss the implications of these distributions for the conservation, cultivation, and ecological restoration of this important species.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EMMAC">
      <title>Material and methods</title>
      <sec sec-type="Study area and species" id="SECID0EQMAC">
        <title>Study area and species</title>
        <p>The study was carried out in West Africa including Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Burkina Faso, Togo, Benin, and Nigeria (Fig. <xref ref-type="fig" rid="F1">1A</xref>). In terms of biogeography, four major phytochoria with specific vegetation types can be broadly distinguished: (i) Sahelian savannah (S), (ii) Sudanian savannah (<abbrev xlink:title="Sudanian savannah" id="ABBRID0E1MAC">SZ</abbrev>), (iii) Guinea savannah (<abbrev xlink:title="Guinea savannah" id="ABBRID0E5MAC">SG</abbrev>), (iv) Guineo-Congolian (<abbrev xlink:title="Guineo-Congolian" id="ABBRID0ECNAC">GC</abbrev>) forest with closed semi-deciduous forests and evergreen forests (Fig. <xref ref-type="fig" rid="F1">1A</xref>). The African lowland rain forest has three regions, namely Congolia, Upper Guinea, and Lower Guinea, the latter two being separated by the Dahomey Gap (<xref ref-type="bibr" rid="B85">White 1983</xref>; <xref ref-type="bibr" rid="B66">Poorter et al. 2004</xref>; <xref ref-type="bibr" rid="B51">Leal 2004</xref>). The current forest range originated from the dry period in the region during the Quaternary, especially the Pleistocene and the Holocene (<xref ref-type="bibr" rid="B69">Salzmann and Hoelzmann 2005</xref>). The Dahomey Gap has been identified as a major ecogeographical impediment to the dispersal of rainforest species in this region and is, as such, of crucial significance for their distribution patterns in West Africa (<xref ref-type="bibr" rid="B82">van Bruggen 1989</xref>; <xref ref-type="bibr" rid="B54">Martin 1991</xref>; <xref ref-type="bibr" rid="B49">Jenik 1994</xref>). The Dahomey Gap experienced a decline in annual precipitation from more than 2000 mm in the rainforest areas of Nigeria and Ivory Coast to about 1200–1000 mm in the savanna region. The <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EGOAC">GC</abbrev> region is relatively dry and receives between 1600 and 2000 mm of precipitation annually, while the <abbrev xlink:title="Sudanian savannah" id="ABBRID0EKOAC">SZ</abbrev> and S zones receive 500–1400 mm and 150–500 mm of annual precipitation respectively.</p>
        <fig id="F1" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure1</object-id>
          <object-id content-type="arpha">D7BDEDA2-E62C-5D3F-ADCA-7E41A5183112</object-id>
          <label>Figure 1.</label>
          <caption>
            <p><bold>A</bold>. Study area with distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> across West Africa (white circles). <bold>B</bold>. Tree. <bold>C</bold>. Leaf. <bold>D</bold>. Fruits. <bold>E</bold>. Suckers. Photographs B–E by G.H. Dassou in 2021.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140425.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140425</uri>
          </graphic>
        </fig>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> is currently present in a wide geographical range across the four ecological zones: <abbrev xlink:title="Guineo-Congolian" id="ABBRID0E4PAC">GC</abbrev>, <abbrev xlink:title="Guinea savannah" id="ABBRID0ECAAE">SG</abbrev>, <abbrev xlink:title="Sudanian savannah" id="ABBRID0EGAAE">SZ</abbrev>, andS (Fig. <xref ref-type="fig" rid="F1">1A</xref>). The species is mainly found in Senegal, Guinea, Sierra Leone, Liberia, Ivory Coast, Ghana, Burkina Faso, Togo, Benin, and Nigeria (<xref ref-type="bibr" rid="B84">WFO 2024</xref>). The species is a tree with a large leafy crown reaching 10–40 m in height (Fig. <xref ref-type="fig" rid="F1">1B</xref>) (<xref ref-type="bibr" rid="B7">Akoègninou et al. 2006</xref>). The trunk is generally greyish and fairly smooth but covered with fine cracks. The wood is dark reddish brown, hard, and fine-grained. The leaves are bright green, pinnate and have 10 alternate leaflets, which are oval to elliptical, thinly coriaceous, or papery, usually 4–6 cm long and 2.5–3 cm wide, with rather few translucid gland-dots (Fig. <xref ref-type="fig" rid="F1">1C</xref>). The inflorescence is a panicle or loose axillary fascicle, dense and lax. The apetalous flowers have a calyx with 4 sepals, creamy white, in bud glabrous or sparsely pubescent outside, and 10 stamens. The ovary is ellipsoid, ca 2 mm long, densely hairy, 1-celled, style 3–4 mm long, arched. The fruit is a globose or subglobose drupe, slightly flattened about 5–6 cm in diameter, thick, fleshy (Fig. <xref ref-type="fig" rid="F1">1D</xref>), and it is divided into three main parts: (i) dark green epicarp, hard for immature fruits, light green tending to brown and brittle for ripe fruits, cracks at maturity, (ii) greenish mesocarp (pulp) intertwined with fibres inserted on the core, (iii) voluminous, woody core covered with fibrous meshes and enclosing a single ovoid and flattened seed of dark brown colour. Flowering occurs from February to May, and fruits take about five to six months to mature. The fruits are dispersed mainly by humans, monkeys, rodents, and parrots (<xref ref-type="bibr" rid="B50">Kouyaté and Lamien 2011</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> also holds a high potential to shoot forth suckers (Fig. <xref ref-type="fig" rid="F1">1E</xref>).</p>
      </sec>
      <sec sec-type="Species occurrence data" id="SECID0EVBAE">
        <title>Species occurrence data</title>
        <p>Occurrence data were gathered from two sources: online repositories and previously published data. Online data included the records from the Global Biodiversity Information Facility (<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://GBIF.org">GBIF.org</ext-link> 2021) and RAINBIO (<xref ref-type="bibr" rid="B20">Dauby et al. 2016</xref>). These occurrences were cleaned by removing duplicates and records without geographic coordinates. In addition, older occurrences collected before 1980 were removed to reduce the effects of temporal bias and match occurrence to the climate datasets (<xref ref-type="bibr" rid="B44">Idohou et al. 2017</xref>). As for previously published data, occurrences were extracted from two papers, namely <xref ref-type="bibr" rid="B17">Dangbo et al. (2019)</xref> and <xref ref-type="bibr" rid="B21">Diop et al. (2010)</xref>. These coordinates were obtained by georeferencing the maps and digitizing the datapoints. To avoid occurrence bias, the global gazetteer v.2.3 (<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://www.fallingrain.com/world">http://www.fallingrain.com/world</ext-link>) was used to assign digital coordinates to the occurrences (<xref ref-type="bibr" rid="B38">Hemami et al. 2020</xref>). We projected these coordinates on Google Earth v.7.1 to ensure they matched the target localities. Finally, all occurrences were combined, and we used the occurrence rarefy tool of Spatial Analyst in ArcGIS v.10.8 to reduce autocorrelation among the occurrences. Thus, we kept only one point within each grid cell (5 × 5 km) to be consistent with the environmental layers. Overall, after the cleaning process, 220 occurrences were used to run the models across West Africa: 175 from GBIF, 36 from RAINBIO, and 9 from the two papers (Supplementary material <xref ref-type="supplementary-material" rid="S1">1</xref>).</p>
      </sec>
      <sec sec-type="Environmental data" id="SECID0E4CAE">
        <title>Environmental data</title>
        <p>A total of three sources of environmental layers (bioclimatic, edaphic, and topography) were used. Past, current, and future climatic data were obtained at the spatial resolution of 2.5 minutes (~5 km at the equator) from WorldClim v.2.1 (<xref ref-type="bibr" rid="B25">Fick and Hijmans 2017</xref>). The edaphic (<xref ref-type="bibr" rid="B52">Leenaars et al. 2014</xref>) whereas elevation variables were taken from the digital elevation model (ASTER DEM) (<xref ref-type="bibr" rid="B25">Fick and Hijmans 2017</xref>). Slope data was generated from Google Earth Engine (<xref ref-type="bibr" rid="B28">Gorelick 2013</xref>) using a West African boundary. The current variables layers were derived from 1950–2000 climate data (<xref ref-type="bibr" rid="B40">Hijmans et al. 2005</xref>). For the past projection, bioclimatic variables layers were obtained from the MIROC-ESM model for the Mid-Holocene (<abbrev xlink:title="Mid-Holocene" id="ABBRID0EXDAE">MH</abbrev>, about 6000 years ago) and the Last Glacial Maximum (<abbrev xlink:title="Last Glacial Maximum" id="ABBRID0E2DAE">LGM</abbrev>, about 22,000 years ago). As for the future projection, the MIROC6 model was used to be consistent with the past projection. MIROC are advanced climate models developed by the Atmosphere and Ocean Research Institute (AORI) of the University of Tokyo (<xref ref-type="bibr" rid="B77">Tatebe et al. 2019</xref>). The benefit of MIROC6 models is that they have improved climate projections (temperature and precipitation), advanced physical processes, and comprehensive earth system modelling (<xref ref-type="bibr" rid="B77">Tatebe et al. 2019</xref>). Past and future layers were also downloaded from the WorldClim website (<xref ref-type="bibr" rid="B25">Fick and Hijmans 2017</xref>). The suitable areas were projected under two climate scenarios, ssp245 and ssp585, for the year 2050 (average for 2041–2060). The downloaded variables cut across WA and were resampled at 5 km. We removed four bioclimatic variables (bio8, bio9, bio18, and bio19) because of discontinuities between neighbouring pixels for the African continent (<xref ref-type="bibr" rid="B55">Montoya-Jiménez et al. 2022</xref>). The bioclimatic data consisted of 15 bioclimatic variables (Supplementary material <xref ref-type="supplementary-material" rid="S2">2</xref>). Then, we performed the Pearson correlation and jackknife tests (Supplementary materials <xref ref-type="supplementary-material" rid="S3">3</xref> and <xref ref-type="supplementary-material" rid="S4">4</xref>) using the R package usdm v.2.1-6 (<xref ref-type="bibr" rid="B56">Naimi 2017</xref>). These two tests were used to exclude highly correlated variables with the coefficient of correlation (r) value of |0.7| and to highlight those with high contribution. The final candidate factors were selected after reviewing the literature to determine which variables were ecologically significant for the species.</p>
      </sec>
      <sec sec-type="Modelling technique" id="SECID0E6EAE">
        <title>Modelling technique</title>
        <p>We modelled the past, current, and future distributions of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in West Africa. Four machine learning methods were used for this purpose, and the results were averaged for the predictive models. These methods included generalized additive models (<abbrev xlink:title="generalized additive models" id="ABBRID0EQFAE">GAM</abbrev>) (<xref ref-type="bibr" rid="B86">Wood 2006</xref>), generalized linear models (<abbrev xlink:title="generalized linear models" id="ABBRID0EYFAE">GLM</abbrev>) (<xref ref-type="bibr" rid="B35">Hastie and Tibshirani 1990</xref>), maximum entropy (<abbrev xlink:title="Maximum Entropy" id="ABBRID0EAGAE">MaxEnt</abbrev>) (<xref ref-type="bibr" rid="B64">Phillips et al. 2006</xref>), and random forest (<abbrev xlink:title="random forest" id="ABBRID0EIGAE">RF</abbrev>) (<xref ref-type="bibr" rid="B13">Breiman 2001</xref>). These models were chosen for their effectiveness and robustness (<xref ref-type="bibr" rid="B67">Rong et al. 2020</xref>; <xref ref-type="bibr" rid="B16">Daï et al. 2023</xref>). The <abbrev xlink:title="random forest" id="ABBRID0EYGAE">RF</abbrev> is a learning technique that constructs several decision trees during training and outputs to the class. <abbrev xlink:title="generalized additive models" id="ABBRID0E3GAE">GAM</abbrev> is a statistical model that is used to link response variables to predictor factors (<xref ref-type="bibr" rid="B86">Wood 2006</xref>). The <abbrev xlink:title="generalized linear models" id="ABBRID0EEHAE">GLM</abbrev> model describes the relationship between the dependent and independent variables (<xref ref-type="bibr" rid="B35">Hastie and Tibshirani 1990</xref>). The <abbrev xlink:title="Maximum Entropy" id="ABBRID0EMHAE">MaxEnt</abbrev> technique is a probabilistic model that aims to find the probability distribution with the highest entropy (or the least uncertainty) given specific restrictions (<xref ref-type="bibr" rid="B64">Phillips et al. 2006</xref>, <xref ref-type="bibr" rid="B65">2017</xref>). All models were run with the R package sdm v.1.1-8 (<xref ref-type="bibr" rid="B57">Naimi and Araújo 2016</xref>). Environmental factors were used as independent variables, whereas point locations (geographical coordinates) were used as dependent variables. We also used 10<sup>3</sup> pseudo-absence points since we do not have the real absence data. Furthermore, the occurrences were split into two samples where 70% were used to train the models and 30% to test them. To increase the models’ accuracy, each model was replicated 10 times with subsampling and bootstrapping. We assessed the accuracy and the performance of the models using the area under the receiver operating characteristic curve (<abbrev xlink:title="operating characteristic curve" id="ABBRID0E5HAE">AUC</abbrev>) and the true skills statistic (<abbrev xlink:title="true skills statistic" id="ABBRID0ECIAE">TSS</abbrev>) (<xref ref-type="bibr" rid="B8">Allouche et al. 2006</xref>). The <abbrev xlink:title="operating characteristic curve" id="ABBRID0EKIAE">AUC</abbrev> provides the probability that the predictive power of a model is better than random prediction (<abbrev xlink:title="operating characteristic curve" id="ABBRID0EOIAE">AUC</abbrev> = 0.5) (<xref ref-type="bibr" rid="B76">Swets 1988</xref>). A model with an <abbrev xlink:title="operating characteristic curve" id="ABBRID0EWIAE">AUC</abbrev> value close to 1 (<abbrev xlink:title="operating characteristic curve" id="ABBRID0E1IAE">AUC</abbrev> ≥ 0.75) is considered to be a good fit. As for <abbrev xlink:title="true skills statistic" id="ABBRID0E5IAE">TSS</abbrev>, values range from -1 to 1 and values close to 1 indicate the high performance of the models.</p>
      </sec>
      <sec sec-type="Range dynamics, direction of range shifts, and conservation status" id="SECID0ECJAE">
        <title>Range dynamics, direction of range shifts, and conservation status</title>
        <p>We evaluated the dynamics of the suitable areas and the direction of range shifts using SDMtoolbox v.1.0b (<xref ref-type="bibr" rid="B14">Brown 2014</xref>). This toolbox is a python-based GIS software that is capable of describing the magnitude and direction of the predicted change over time by summarizing the core shifts in the species distribution reducing the distribution variation to a single centroid (central) point and generating a vector file (<xref ref-type="bibr" rid="B14">Brown 2014</xref>). In our case, the range shifts, as well as range dynamics, were assessed by comparing Mid-Holocene (<abbrev xlink:title="Mid-Holocene" id="ABBRID0EQJAE">MH</abbrev>)-current, Last Glacial Maximum (<abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EUJAE">LGM</abbrev>)-current, current-ssp245, and current-ssp585. We also evaluated the dynamics of the suitable areas using Distribution Changes Between Binary SDMs (<xref ref-type="bibr" rid="B14">Brown 2014</xref>). The process was the same as described above but here the tool generated a single layer in TIF format and an Excel sheet that held the statistics. Overall, four classes of habitats were automatically generated: contraction, expansion, areas of no change or stability, and no occupancy areas or unsuitable.</p>
        <p>The conservation status of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> was evaluated based on the IUCN criteria, especially B. We determined the extent of occurrence (<abbrev xlink:title="extent of occurrence" id="ABBRID0EJKAE">EOO</abbrev>) and the area of occupancy (<abbrev xlink:title="area of occupancy" id="ABBRID0ENKAE">AOO</abbrev>) with the R package ConR v.1.3.0 (<xref ref-type="bibr" rid="B19">Dauby and Lima 2023</xref>). The <abbrev xlink:title="extent of occurrence" id="ABBRID0EVKAE">EOO</abbrev> is a polygon that can be drawn to encompass all the known present occurrences of a species (<xref ref-type="bibr" rid="B47">IUCN 2012</xref>). The <abbrev xlink:title="area of occupancy" id="ABBRID0E4KAE">AOO</abbrev> is related to the area covered by the number of occupied cells of a grid for a given resolution (<xref ref-type="bibr" rid="B47">IUCN 2012</xref>). In this study, we considered grid cells of 2 km<sup>2</sup>, as recommended by IUCN guidelines (<xref ref-type="bibr" rid="B48">IUCN 2024</xref>). In addition, the diverse threat levels were measured by estimating the number of locations in protected areas. For this purpose, the occurrence of the species within the West African boundary (<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="https://geoportal.icpac.net">https://geoportal.icpac.net</ext-link>) and within protected areas (World Database on Protected Areas, <xref ref-type="bibr" rid="B81">UNEP-WCMC 2023</xref>) was used.</p>
      </sec>
    </sec>
    <sec sec-type="Results" id="SECID0EULAE">
      <title>Results</title>
      <sec sec-type="Factors affecting D. senegalense distribution and model accuracy" id="SECID0EYLAE">
        <title>Factors affecting <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution and model accuracy</title>
        <p>Overall, five variables were identified to influence the distribution range of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> (Fig. <xref ref-type="fig" rid="F2">2A</xref>). These variables were related to temperature and precipitation. Isothermality (bio3) was the variable that contributed the most (41.10%). The occurrence probability of the species reached its maximal value when isothermality values were close to 100 (Supplementary material <xref ref-type="supplementary-material" rid="S5">5</xref>). This variable was followed by the Precipitation of Wettest Month (bio13, 21.50%) and the Precipitation Seasonality (bio15, 19.23%). For bio13, the occurrence probability of the species increased when the bio13 values increased. In contrast, species occurrence decreased when the bio15 values varied by around 35%. The Temperature Annual Range (bio7, 10.60%) decreased the occurrence probability of the species with bio7 values below 75 mm (Supplementary material <xref ref-type="supplementary-material" rid="S5">5</xref>). Even though bio7 had a minor impact on species distribution, the occurrence probability of the species tended to increase between values of 10°C and 40°C (Supplementary material <xref ref-type="supplementary-material" rid="S5">5</xref>). Precipitation of Driest Month (bio14) showed the lowest contribution (7.57%). The performance and accuracy of the models showed an average area under the curve (<abbrev xlink:title="operating characteristic curve" id="ABBRID0EFNAE">AUC</abbrev>) of 0.92 and a <abbrev xlink:title="true skills statistic" id="ABBRID0EJNAE">TSS</abbrev> value of 0.73 (Fig. <xref ref-type="fig" rid="F2">2B</xref>), indicating that the models are of excellent quality.</p>
        <fig id="F2" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure2</object-id>
          <object-id content-type="arpha">FD8797B4-F324-5C63-99D2-F468787CCF5D</object-id>
          <label>Figure 2.</label>
          <caption>
            <p>Diagram showing the contribution of the main variables and the performance of the model. <bold>A</bold>. Contribution of the variables involved in the model. Each variable is represented by a code: bio3 = Isothermality, bio7 = Temperature Annual Range, bio13 = Precipitation of Wettest Month, bio14 = Precipitation of Driest Month, bio15 = Precipitation Seasonality. <bold>B</bold>. Performance of the four algorithms. Values in the y-axis represent the average of 10 repetitions related to the two metrics. <abbrev xlink:title="operating characteristic curve" id="ABBRID0E4NAE">AUC</abbrev> = Area Under the Curve and <abbrev xlink:title="true skills statistic" id="ABBRID0EBOAE">TSS</abbrev> = True Skill Statistic.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140426.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140426</uri>
          </graphic>
        </fig>
      </sec>
      <sec sec-type="Potential suitability distribution of D. senegalense under past, current, and future climatic conditions" id="SECID0EKOAE">
        <title>Potential suitability distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> under past, current, and future climatic conditions</title>
        <p>The extent of suitability shown for the four classes (expansion, unsuitable, stable, contraction) for the five time periods (<abbrev xlink:title="Last Glacial Maximum" id="ABBRID0E3OAE">LGM</abbrev>, <abbrev xlink:title="Mid-Holocene" id="ABBRID0EAPAE">MH</abbrev>, current, ssp245, ssp585) is documented in Supplementary material <xref ref-type="supplementary-material" rid="S6">6</xref>. In the past, notably at the <abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EIPAE">LGM</abbrev>, the stable ranges of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution, estimated at 1,183,095.65 km<sup>2</sup> or 21.30% of the study area, mainly occurred from Nigeria to Sierra Leone, and from the Guinean savannah to the evergreen forest ecological zones (Fig. <xref ref-type="fig" rid="F3">3A</xref>). Overall, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> was found across the <abbrev xlink:title="Sudanian savannah" id="ABBRID0EIQAE">SZ</abbrev> to <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EMQAE">GC</abbrev> zones of Nigeria and Benin, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EQQAE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EUQAE">GC</abbrev> zones of Togo, and from the <abbrev xlink:title="Guinea savannah" id="ABBRID0EYQAE">SG</abbrev> to the evergreen forest zone of Ivory Coast. The species was also found from the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0E3QAE">GC</abbrev> to the evergreen forest zone of Guinea and Sierra Leone and the evergreen forest zone of Liberia. A very slight range expansion of 241,458.47 km<sup>2</sup> or 0.16% was observed during this period, mainly in the extreme west of Guinea. Meanwhile, the main part of the study area (3,803,962.10 km<sup>2</sup> or 66.15%) was unsuitable (Fig. <xref ref-type="fig" rid="F3">3A, F</xref>). The species lost an extent of 807,233.78 km<sup>2</sup> or 12.39% from the <abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EKRAE">LGM</abbrev> to the current day.</p>
        <fig id="F3" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure3</object-id>
          <object-id content-type="arpha">85453A44-F99F-5158-8A02-962D0978418B</object-id>
          <label>Figure 3.</label>
          <caption>
            <p>Dynamics of habitat suitability for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> across West Africa. <bold>A</bold>. Last Glacial Maximum (about 22,000 years ago). <bold>B</bold>. Mid-Holocene (about 6000 years ago). <bold>C</bold>. Current distribution. <bold>D</bold>. Future distribution ssp245-2050. <bold>E</bold>. Future distribution ssp585-2050. <bold>F</bold>. Range suitability.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140427.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140427</uri>
          </graphic>
        </fig>
        <p>The <abbrev xlink:title="Mid-Holocene" id="ABBRID0EUSAE">MH</abbrev> period exhibited a constant favourable range of 1,362,492.32 km<sup>2</sup> or 21.50% with an unsuitable range of 3,885,769.34 km<sup>2</sup> or 60.10% (Fig. <xref ref-type="fig" rid="F3">3B</xref>). If we compare the <abbrev xlink:title="Mid-Holocene" id="ABBRID0EATAE">MH</abbrev> to the current period, there was no discernible rise in the stable distribution range. The increase was estimated at 62,067.80 km<sup>2</sup> or 0.03% of the total study area (Fig. <xref ref-type="fig" rid="F3">3B, F</xref>). The contraction range of the species’ distribution area was estimated at 725,420.54 km<sup>2</sup> or 18.37%.</p>
        <p>At present, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> occurs in a large geographical range across Senegal, Guinea, Guinea-Bissau, Sierra Leone, Liberia, Gambia, Ivory Coast, Ghana, Burkina Faso, Togo, Benin, and Nigeria, and spreading across four ecological zones: Guineo-Congolian (<abbrev xlink:title="Guineo-Congolian" id="ABBRID0EZTAE">GC</abbrev>), Guinea savannah (<abbrev xlink:title="Guinea savannah" id="ABBRID0E4TAE">SG</abbrev>), Sudanian savannah (<abbrev xlink:title="Sudanian savannah" id="ABBRID0EBUAE">SZ</abbrev>), and very lower part of the Sahelian savannah (S) zones (Fig. <xref ref-type="fig" rid="F3">3C</xref>). The species occupies an extent of 1,048,558.40 km<sup>2</sup> in West Africa. These suitable areas or stable areas for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution under current climatic conditions are located between 5°00’00”N–11°00’00”N and 10°00’00”W–5°00’00”W in the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EWUAE">GC</abbrev> and <abbrev xlink:title="Guinea savannah" id="ABBRID0E1UAE">SG</abbrev> zones (Fig. <xref ref-type="fig" rid="F3">3C</xref>). Countries such as Ivory Coast, Nigeria, Benin, Togo, Ghana, Liberia, and Sierra Leone showed the largest stable ranges. The countries where <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> was less represented are Burkina-Faso, Guinea-Bissau, Guinea, Gambia, and Senegal. Overall, 3,954,573.16 km<sup>2</sup> of the total West Africa was not suitable for the species under current conditions.</p>
        <p>Regarding the potential distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in the future, the stable ranges were 506,147.78 km<sup>2</sup> or 17.70% under ssp245 and 157,558 km<sup>2</sup> or 13.98% under ssp585 by horizon-time 2050 (Fig. <xref ref-type="fig" rid="F3">3D, E</xref>). Results also showed that the unsuitable range predicted for the future appeared to be higher in both scenarios (ssp245 and ssp585) than in the <abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EEWAE">LGM</abbrev> and <abbrev xlink:title="Mid-Holocene" id="ABBRID0EIWAE">MH</abbrev> periods respectively. These unsuitable areas were 5,342,985.84 km<sup>2</sup> or 78.70% and 5,551,460.57 km<sup>2</sup> or 78.13% under ssp245 and ssp585 respectively. However, these two scenarios showed much smaller contraction ranges than the two previous periods (<abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EQWAE">LGM</abbrev> and <abbrev xlink:title="Mid-Holocene" id="ABBRID0EUWAE">MH</abbrev>), with 181,355.14 km<sup>2</sup> or 2.93% for the ssp245 scenario and 322,893.51 km<sup>2</sup> or 7.29% for the ssp585 scenario. At the same time, some slight expansions occurred estimating around 5,261.25 km<sup>2</sup> or 0.67% for ssp245 and 3,837.92 km<sup>2</sup> or 0.60% for ssp585. In terms of spatial distribution, suitable areas are expected in the <abbrev xlink:title="Sudanian savannah" id="ABBRID0EAXAE">SZ</abbrev> to the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EEXAE">GC</abbrev> zones of Nigeria and Benin, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EIXAE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EMXAE">GC</abbrev> zones of Togo, and from the <abbrev xlink:title="Guinea savannah" id="ABBRID0EQXAE">SG</abbrev> to the evergreen forest zone of Ivory Coast. However, the species’ suitable areas have shrunk on both sides of the Ivory Coast-Ghana border, with a more pronounced loss in the ssp58 scenario.</p>
      </sec>
      <sec sec-type="Distributional shifts of the centroid of D. senegalense in West Africa" id="SECID0EUXAE">
        <title>Distributional shifts of the centroid of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in West Africa</title>
        <p>The centroid of the suitable area for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> revealed different patterns of range change shift direction (Fig. <xref ref-type="fig" rid="F4">4</xref>). In past, especially during <abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EVYAE">LGM</abbrev>, range shift directions tended to be more longitudinal rather than latitudinal, with a north-eastern shift for <abbrev xlink:title="Last Glacial Maximum" id="ABBRID0EZYAE">LGM</abbrev> and a south-western shift for <abbrev xlink:title="Mid-Holocene" id="ABBRID0E4YAE">MH</abbrev>. However, both future scenarios showed range shifts to the south-east (Fig. <xref ref-type="fig" rid="F4">4</xref>). The current centre of distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> was predicted to be in the extreme east of Ivory Coast (Fig. <xref ref-type="fig" rid="F4">4</xref>).</p>
        <fig id="F4" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure4</object-id>
          <object-id content-type="arpha">3C7917F9-F94D-570A-85D6-DD66B664182B</object-id>
          <label>Figure 4.</label>
          <caption>
            <p>Range shift direction under past (blue and red arrows) and future scenarios (yellow and purple arrows). S: Sahelian Savannah, <abbrev xlink:title="Sudanian savannah" id="ABBRID0E3ZAE">SZ</abbrev>: Sudanian Savannah, <abbrev xlink:title="Guinea savannah" id="ABBRID0EA1AE">SG</abbrev>: Guinea Savannah, <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EE1AE">GC</abbrev>: Guineo-Congolian.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140428.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140428</uri>
          </graphic>
        </fig>
      </sec>
      <sec sec-type="Distribution analysis of D. senegalense within protected areas (PAs) and its conservation assessment" id="SECID0EN1AE">
        <title>Distribution analysis of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> within protected areas (PAs) and its conservation assessment</title>
        <p>All PAs located between 6°N–13°N and 12°W–10°E were currently suitable for the species (Fig. <xref ref-type="fig" rid="F5">5</xref>). The models indicated that all PAs in Ivory Coast and Liberia are suitable for the species. In Benin, the PAs suitable for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> were those in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EO2AE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0ES2AE">GC</abbrev> zones. There were no suitable areas in the <abbrev xlink:title="Sudanian savannah" id="ABBRID0EW2AE">SZ</abbrev>, including the Pendjari and W parks. Similar observations showed that the PAs in Nigeria, Togo, and Ghana that are currently suitable for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> were located in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EF3AE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EJ3AE">GC</abbrev> zones, but more predominantly in the central east and south of the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EN3AE">GC</abbrev> zone and in the west of the <abbrev xlink:title="Guinea savannah" id="ABBRID0ER3AE">SG</abbrev> zone of Nigeria, in the south-east of the <abbrev xlink:title="Guinea savannah" id="ABBRID0EV3AE">SG</abbrev> zone and throughout the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EZ3AE">GC</abbrev> zone of Ghana, and from halfway the <abbrev xlink:title="Guinea savannah" id="ABBRID0E43AE">SG</abbrev> zone to the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EB4AE">GC</abbrev> zone of Togo. In Guinea, the suitable PAs were located in the south-east of the country in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EF4AE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EJ4AE">GC</abbrev> zones. Only the PAs along the coast of Senegal, Guinea-Bissau, and Gambia were capable of hosting the species in the present day. In addition, the PAs in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EN4AE">SG</abbrev> zone of Burkina Faso were suitable for the species under current conditions.</p>
        <fig id="F5" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure5</object-id>
          <object-id content-type="arpha">CEF37C12-0799-5242-8EF9-97F8139F9103</object-id>
          <label>Figure 5.</label>
          <caption>
            <p>Spatial distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in protected areas under current and future scenarios.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140430.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140430</uri>
          </graphic>
        </fig>
        <p>According to the future scenario (ssp245), most of the protected areas that are suitable under current conditions should also be suitable for the species by 2050. However, some unsuitable protected areas are predicted under this scenario. These are located mainly in the south-west and north-west of Ghana and the south-east and north of Ivory Coast, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EL5AE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EP5AE">GC</abbrev> zones respectively. In addition, some unsuitable PAs are predicted to a slight extent in the <abbrev xlink:title="Sudanian savannah" id="ABBRID0ET5AE">SZ</abbrev> zone to the east of the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EX5AE">GC</abbrev> zone in Nigeria and in the east of the <abbrev xlink:title="Guinea savannah" id="ABBRID0E25AE">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0E65AE">GC</abbrev> zones in Guinea. The extent of suitability related to each class within the protected areas is provided in Supplementary material <xref ref-type="supplementary-material" rid="S6">6</xref>. The number of protected areas by country as well as their extent were also quantified. Overall, the models indicated that in the present conditions, only 202,569.67 km<sup>2</sup> or 22.90% of the 2011 PAs of West Africa are suitable for the species. In contrast, 698,554.08 km<sup>2</sup> or 77.10% of the total extent (i.e. 901,123.75 km<sup>2</sup>) is not suitable. Moreover, 98,915.92 km<sup>2</sup> or 21.01% and 27,573.05 km<sup>2</sup> or 14.60% could be suitable under ssp245 and ssp585 by 2050 respectively. Significant contraction was observed under the ssp585 scenario with 21,7141.24 km<sup>2</sup> or 8.61%. The contraction predicted under ssp585 was 145,798.37 km<sup>2</sup> or 2.50%. Suitable areas or stable habitats were found especially in Liberia, central and southern Sierra Leone, in the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EV6AE">GC</abbrev> zone of southern Nigeria, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EZ6AE">SG</abbrev> zone of Benin and Togo and the south-west of Ivory Coast, and in a few protected areas in central and south-eastern Ghana. The expansion of habitat was estimated at around 413.18 km<sup>2</sup> or 0.51% and 468.27 km<sup>2</sup> or 0.35% under ssp245 and ssp585 respectively. As for unsuitable areas, the estimations were 698,554.08 km<sup>2</sup> or 77.10% under current conditions. Future scenarios were also high with 655,996.27 km<sup>2</sup> or 75.98% for ssp245 and 655,941.18 km<sup>2</sup> or 76.44% under ssp585 by 2050.</p>
        <p>The status of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> based on IUCN criteria across West Africa was documented in Fig. <xref ref-type="fig" rid="F6">6</xref>. The results showed an <abbrev xlink:title="extent of occurrence" id="ABBRID0EZAAG">EOO</abbrev> of 1,958,696 km<sup>2</sup> (&gt; 20,000 km<sup>2</sup>) and an <abbrev xlink:title="area of occupancy" id="ABBRID0EBBAG">AOO</abbrev> of 668 km<sup>2</sup> (&lt; 2,000 km<sup>2</sup>). The large <abbrev xlink:title="extent of occurrence" id="ABBRID0EJBAG">EOO</abbrev> points towards Least Concern, while the <abbrev xlink:title="area of occupancy" id="ABBRID0ENBAG">AOO</abbrev> could indicate Vulnerable (VU). However, the results also indicated that 30.8% of the 220 unique occurrences are located within 44 protected areas. Across the study area, we found 130 subpopulations (considering a 2 km<sup>2</sup> radius) and 151 locations (considering a grid size of 10 km<sup>2</sup>). Therefore, the species can be considered as Least Concern in West Africa.</p>
        <fig id="F6" position="float" orientation="portrait">
          <object-id content-type="doi">10.5091/plecevo.122470.figure6</object-id>
          <object-id content-type="arpha">40AB84FD-B4DE-5E21-8F24-93CC48BB9E0E</object-id>
          <label>Figure 6.</label>
          <caption>
            <p>Conservation assessement of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> based on IUCN criteria across West Africa.</p>
          </caption>
          <graphic xlink:href="plecevo-157-343-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1140431.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1140431</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="SECID0ENCAG">
      <title>Discussion</title>
      <sec sec-type="Importance of climatic factors on species distribution and model performance" id="SECID0ERCAG">
        <title>Importance of climatic factors on species distribution and model performance</title>
        <p>Environmental factors play an important role in forest establishment and persistence (<xref ref-type="bibr" rid="B29">Greve et al. 2011</xref>). As a result, under climate change, a set of factors that favour the distribution of a given plant species, particularly multipurpose trees such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic>, may help elucidate its resilience. In this study, climatic factors (temperature and precipitation) were the major variables that explained the species distribution in West Africa consistent with the ecological requirements listed by <xref ref-type="bibr" rid="B21">Diop et al. (2010)</xref>. However, this finding differs from that of <xref ref-type="bibr" rid="B5">Agbo et al. (2019)</xref> who found that soil factors (cation exchange capacity) were important for the distribution of the related species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="microcarpum">microcarpum</tp:taxon-name-part></tp:taxon-name></italic> in Benin. Even though soil properties are recommended in plant modelling (<xref ref-type="bibr" rid="B39">Hengl et al. 2017</xref>; <xref ref-type="bibr" rid="B87">Zuquim et al. 2020</xref>), their incorporation is meaningful in local contexts in contrast to regional ones, as is the case in this study (<xref ref-type="bibr" rid="B62">Pearson and Dawson 2003</xref>).</p>
        <p>The distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> throughout West Africa was mainly affected by isothermality (bio3). Overall, isothermality refers to how similar the average daily maximum and minimum temperatures are throughout the year (<xref ref-type="bibr" rid="B61">O’Donnel and Ignizio 2012</xref>). Here, values were close to 100, indicating that the species preferred areas with consistent diurnal temperatures throughout the year. This factor is a determinant of plant distribution in tropical areas including West Africa (<xref ref-type="bibr" rid="B43">Huang et al. 2021</xref>). In natural ecosystems, the species is primarily distributed along rivers in gallery forests with an annual precipitation of 800 to 1800 mm, in contrast to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="microcarpum">microcarpum</tp:taxon-name-part></tp:taxon-name></italic>, which is more commonly found in open savannah (<xref ref-type="bibr" rid="B17">Dangbo et al. 2019</xref>; <xref ref-type="bibr" rid="B41">Houénon et al. 2022</xref>). Our findings also showed that species occurrence increased when the precipitation during the wettest month (bio13) increased, which explains why species are more common in rainy ecosystems. In contrast, species occurrence probability decreased when the precipitation of the driest month (bio14) was around 75 mm. Similarly, the occurrence probability decreased for precipitation seasonality (bio15) values of around 35%. This explains how a slight variation in precipitation in a season, particularly during the dry seasons, could influence species seed dispersal, which is facilitated by water and animals. The annual temperature range (bio7) had a minor impact on species distribution, but the occurrence probability of the species increased for temperatures between 10°C and 40°C. These findings suggested that the species may have adapted to the wide range of temperatures under climate change but more research is needed to understand how the species might respond to drought, including germination rates, growth rates, and overall plant health. Moreover, the model performance revealed an <abbrev xlink:title="operating characteristic curve" id="ABBRID0ENFAG">AUC</abbrev> value of 0.92, indicating the robustness of the models (<xref ref-type="bibr" rid="B44">Idohou et al. 2017</xref>; <xref ref-type="bibr" rid="B5">Agbo et al. 2019</xref>). Therefore, they could be considered steady and adequate for constructing the potential ecological niche and conservation ranges of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> through its geographical distribution (<xref ref-type="bibr" rid="B24">Favi et al. 2022</xref>).</p>
      </sec>
      <sec sec-type="Impact of climate on species distribution" id="SECID0EIGAG">
        <title>Impact of climate on species distribution</title>
        <p>The current study assessed the dynamics of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution from the past to the predicted future suitable areas. In general, our findings revealed that the suitability range was relatively stable from the last glacial maximum to the present day. In addition, suitable habitats occurred from Nigeria to Sierra Leone, and from the Guinean savannah to the evergreen forest ecological zones. However, a study of a population in Togo indicated that adult individuals are rare and that there is a low regeneration rate, and thus a predominance of small-diameter trees (<xref ref-type="bibr" rid="B17">Dangbo et al. 2019</xref>). Similar results were recently found in Benin by <xref ref-type="bibr" rid="B41">Houénon et al. (2022)</xref> and this shows that modelling results need to be considered with caution. Indeed, human pressures on tree species in the Sub-Saharan region, such as illegal cutting and agricultural expansion, have increased ecosystem vulnerability to climate change, affecting their spatial distribution (<xref ref-type="bibr" rid="B30">Gross et al. 2018</xref>).</p>
        <p>Moreover, the suitable areas of the species shifted from low latitudes to high latitudes from the Last Glacial Maximum to the Mid-Holocene. These findings are consistent with those of <xref ref-type="bibr" rid="B71">Saupe et al. (2019)</xref> about the high-latitudinal migration of tropical species as temperatures rise over geological ages. The Dahomey-gap, which was established during the Holocene, created a unique climate condition in the Guinean Gulf, allowing plant species to spread across a variety of climate zones (<xref ref-type="bibr" rid="B69">Salzmann and Hoelzmann 2005</xref>).</p>
        <p>According to <xref ref-type="bibr" rid="B80">Trisos et al. (2022)</xref>, West Africa is one of the most vulnerable zones to climate change. Since the Industrial Revolution, the average annual temperature has risen above 1.1°C, affecting already the growth and productivity of plant species. Global average annual temperature is expected to reach 5°C by 2100 under worse scenarios (<xref ref-type="bibr" rid="B78">Tokarska et al. 2020</xref>; <xref ref-type="bibr" rid="B15">Chai et al. 2021</xref>). In this context, the models predicted a moderate range decrease for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic>, especially under the pessimistic scenario ssp585. In terms of spatial distribution, suitable areas are expected in the <abbrev xlink:title="Sudanian savannah" id="ABBRID0EIIAG">SZ</abbrev> to <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EMIAG">GC</abbrev> zones of Nigeria and Benin, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EQIAG">SG</abbrev> and <abbrev xlink:title="Guineo-Congolian" id="ABBRID0EUIAG">GC</abbrev> zones of Togo, and from the <abbrev xlink:title="Guinea savannah" id="ABBRID0EYIAG">SG</abbrev> to the evergreen forest zone of Ivory Coast. Previous studies predicted that the congeneric species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="microcarpum">microcarpum</tp:taxon-name-part></tp:taxon-name></italic> could expand in suitable areas (<xref ref-type="bibr" rid="B5">Agbo et al. 2019</xref>). Indeed, co-occurring species may react differently in the same envelop spaces, contracting or expanding their suitable areas in response to global climate change (<xref ref-type="bibr" rid="B33">Hällfors et al. 2016</xref>). This could be supported by the acclimatization or phenotypic plasticity of species to escape extinction (<xref ref-type="bibr" rid="B73">Silva et al. 2019</xref>). Given the increasing warming climate conditions predicted in Africa, most native plant species will experience a decline in the future, e.g. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parkia">Parkia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="biglobosa">biglobosa</tp:taxon-name-part></tp:taxon-name></italic> (Jacq.) R.Br. ex G.Don (<xref ref-type="bibr" rid="B10">Ayihouenou et al. 2016</xref>), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Borassus">Borassus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="aethiopum">aethiopum</tp:taxon-name-part></tp:taxon-name></italic> Mart. (<xref ref-type="bibr" rid="B68">Salako et al. 2019</xref>). Therefore, the hypothesis of a warming climate inducing a decline in the distribution range of a plant species seems to be well-supported (<xref ref-type="bibr" rid="B53">Li et al. 2019</xref>; <xref ref-type="bibr" rid="B45">Imorou 2020</xref>). Moreover, suitable habitat tends to shift to the south-east (wettest areas or low latitudes) in response to future climate scenarios. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">Detarium</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> usually occurs in West African riparian forests (<xref ref-type="bibr" rid="B9">Arbonnier 2009</xref>), from Upper and Lower Guinea to the Dahomey Gap, and in Central Africa (<xref ref-type="bibr" rid="B7">Akoègninou et al. 2006</xref>; <xref ref-type="bibr" rid="B84">WFO 2024</xref>). This region of the continent has a climate with an average annual precipitation of around 1,800 mm, which is ideal for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic>, which has moderate water requirements.</p>
      </sec>
      <sec sec-type="Conservation status, implications for cultivation, ecological restoration and efficient conservation, and model limitations" id="SECID0E2LAG">
        <title>Conservation status, implications for cultivation, ecological restoration and efficient conservation, and model limitations</title>
        <p>Ecological niche modelling or species niche modelling is a powerful and widely used tool to predict the occurrence of many taxa based on environmental factors (<xref ref-type="bibr" rid="B31">Guisan and Zimmermann 2000</xref>). In this study, ENMs were used to have a clear picture of the past, present, and future distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic>.</p>
        <p>The study also addressed the conservation status of the species under current conditions. Our models predicted the suitable areas for the conservation of the target species were located in the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0ESMAG">GC</abbrev> zones of Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, and Nigeria. Furthermore, most protected areas that are suitable under current conditions should be suitable for the species by 2050, except for the south-west and north-west of Ghana and the south-east and north of Ivory Coast, in the <abbrev xlink:title="Guinea savannah" id="ABBRID0EWMAG">SG</abbrev> and Guinean climatic zones, respectively. Based on these findings, we recommend both in situ and ex situ conservation, as the species’ suitable habitats are expected to decrease both within and outside of protected areas. According to <xref ref-type="bibr" rid="B11">BGCI and IUCN (2019)</xref>, the species is of Least Concern. In addition to this statement, they indicated that there are no significant current or future threats to this species. Our findings also classify the species as Least Concern. However, only 30.8% of individuals were found within protected areas. The most important populations are distributed outside protected areas where they are affected by the encroachment due to the agriculture expansion and tree cutting (<xref ref-type="bibr" rid="B18">Dassou et al. 2023</xref>). Previous studies also revealed that the commercial exploitation of timber and medicinal products, associated with the growing regional trade of fruits and seeds, and a low potential for natural regeneration of seeds constitute significant threats to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B22">El-Kamali 2011</xref>; <xref ref-type="bibr" rid="B42">Houénon et al. 2021</xref>, <xref ref-type="bibr" rid="B41">2022</xref>). Nowadays, traditional agroforestry systems are another way to preserve tree species within agricultural lands (<xref ref-type="bibr" rid="B72">Sharma et al. 2016</xref>). It is also possible to introduce the species in well-protected community and sacred forests in the <abbrev xlink:title="Guineo-Congolian" id="ABBRID0E4NAG">GC</abbrev> zone. However, for conservation to be effective, greenhouse experiments must be conducted first, as the species has a low natural regeneration. The outputs of these experiments will be a mainstay for the publication of technical manuals for nursery management. The vision would be to share these techniques with various conservation organisations (NGOs, forestry departments) to facilitate seedling production for the National Days of the Tree in its distribution range and to promote the installation of plantations of the species. Implementing these participatory management policies across local communities and introducing them into agroforestry systems will safely foster species sustainability (<xref ref-type="bibr" rid="B63">Peters and Simaens 2020</xref>). The introduction of the species into the botanical gardens must be also envisaged.</p>
        <p>Given the usefulness of species distribution models in conservation research (<xref ref-type="bibr" rid="B75">Stockwell and Peterson 2002</xref>), the findings of this study are important since they improve insights into the potential changes in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> distribution. The success of model predictions in this study was likely due to the sample size and wider range of data collected, reducing inaccurate predictions caused by incomplete data (<xref ref-type="bibr" rid="B83">Wang et al. 2021</xref>). Even though the models showed interesting performance, models can be improved including physiological data, seed dispersal capacity of the species, and local adaptation data (<xref ref-type="bibr" rid="B23">Engler et al. 2012</xref>; <xref ref-type="bibr" rid="B4">Adoukonou-Sagbadja et al. 2023</xref>). Recently, some new approaches have emerged incorporating intraspecific variation data into the models. For instance, in Benin, the response of the population of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pterocarpus">Pterocarpus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="erinaceus">erinaceus</tp:taxon-name-part></tp:taxon-name></italic> Poir. to climate change was assessed using this method (<xref ref-type="bibr" rid="B12">Biaou et al. 2023</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="Conclusion" id="SECID0ERPAG">
      <title>Conclusion</title>
      <p>By reducing areas with suitable climatic conditions, climate change may disrupt key ecological interactions and compromise species sustainability and the dynamics of populations of plants in their geographical ranges. This study revealed that the current suitable environmental areas for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> would contract under future climate change scenarios across West Africa. The increasing unsuitability was also predicted in protected areas, pointing to a need for setting up sustainable management plans. Isothermality and precipitation were shown to be the main factors that govern the distribution of the species and provided insight into their ecological requirements. Given the announced decline in habitat suitability of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Detarium">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="senegalense">senegalense</tp:taxon-name-part></tp:taxon-name></italic> in the future, associated with threats due to overuse for seed trading, timber, and medicinal products, and the low regeneration potential of this species, in situ conservation may not be a one-stop solution but should be complemented with ex situ conservation. More broadly, the findings of this study could lead to consistent knowledge on the future potential distribution of West African species and appear fundamental to reducing species loss through early sustainable management strategies.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We thank the International Foundation for Science for supporting the work through Grant N° I-1-D-6362 given to Gbèwonmèdéa Hospice Dassou. The authors extend their appreciation to the editor and reviewers for their useful comments and suggestions on the manuscript.</p>
    </ack>
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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl1</object-id>
        <object-id content-type="arpha">D01AE363-B63B-515A-8530-16F4BFF14A51</object-id>
        <label>Supplementary material 1</label>
        <caption>
          <p>Occurrences dataset.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s001.csv" mimetype="text" mime-subtype="csv" position="float" orientation="portrait" xlink:type="simple" id="oo_1140432.csv">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140432</uri>
        </media>
      </supplementary-material>
      <supplementary-material id="S2" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl2</object-id>
        <object-id content-type="arpha">E86AC08D-7A0B-544C-94DF-5F97B751B0C0</object-id>
        <label>Supplementary material 2</label>
        <caption>
          <p>Description of the environmental variables.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s002.csv" mimetype="text" mime-subtype="csv" position="float" orientation="portrait" xlink:type="simple" id="oo_1140433.csv">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140433</uri>
        </media>
      </supplementary-material>
      <supplementary-material id="S3" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl3</object-id>
        <object-id content-type="arpha">E531D054-9130-5367-A2EF-EC27E145F255</object-id>
        <label>Supplementary material 3</label>
        <caption>
          <p>Remaining variables after the Pearson Correlation test.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s003.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" xlink:type="simple" id="oo_1140434.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140434</uri>
        </media>
      </supplementary-material>
      <supplementary-material id="S4" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl4</object-id>
        <object-id content-type="arpha">2E466599-CD0F-50DC-AB9D-B03D7189B826</object-id>
        <label>Supplementary material 4</label>
        <caption>
          <p>Variables contributions obtained from the jackknife test.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s004.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" xlink:type="simple" id="oo_1140435.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140435</uri>
        </media>
      </supplementary-material>
      <supplementary-material id="S5" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl5</object-id>
        <object-id content-type="arpha">611E8C47-E39B-5158-8D60-81D6137BCA2E</object-id>
        <label>Supplementary material 5</label>
        <caption>
          <p>Average of the response curves for the four models used.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s005.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" xlink:type="simple" id="oo_1140436.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140436</uri>
        </media>
      </supplementary-material>
      <supplementary-material id="S6" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.5091/plecevo.122470.suppl6</object-id>
        <object-id content-type="arpha">6B66A67E-7501-5EC1-9D41-2E3286A3EF60</object-id>
        <label>Supplementary material 6</label>
        <caption>
          <p>Estimations of suitable area in km<sup>2</sup> for the whole study area and protected areas.</p>
        </caption>
        <media xlink:href="plecevo-157-343-s006.csv" mimetype="text" mime-subtype="csv" position="float" orientation="portrait" xlink:type="simple" id="oo_1140437.csv">
          <uri content-type="original_file">https://binary.pensoft.net/file/1140437</uri>
        </media>
      </supplementary-material>
    </sec>
  </back>
</article>
