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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.175775</article-id>
      <article-id pub-id-type="publisher-id">175775</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>Core Eudicots: Asterids</subject>
          <subject>Gentianales</subject>
          <subject>Rubiaceae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biodiversity &amp; Conservation</subject>
          <subject>Conservation Biology</subject>
          <subject>Molecular systematics</subject>
          <subject>Phylogeny</subject>
          <subject>Systematics</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Americas</subject>
          <subject>Peru</subject>
          <subject>South America</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name>), a new species with an isolated phylogenetic position and distinct morphology from northern Peru</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Montesinos-Tubée</surname>
            <given-names>Daniel B.</given-names>
          </name>
          <email xlink:type="simple">dmontesinos@cientifica.edu.pe</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-4439-5089</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-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/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>Dasher</surname>
            <given-names>Cameron E.L.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-8329-2149</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <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>
          <role content-type="http://credit.niso.org/contributor-roles/visualization/">Visualization</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Borsch</surname>
            <given-names>Thomas</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-5724-2786</uri>
          <xref ref-type="aff" rid="A2">2</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/formal-analysis/">Formal analysis</role>
          <role content-type="http://credit.niso.org/contributor-roles/funding-acquisition/">Funding acquisition</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>
          <role content-type="http://credit.niso.org/contributor-roles/resources/">Resources</role>
          <role content-type="http://credit.niso.org/contributor-roles/software/">Software</role>
          <role content-type="http://credit.niso.org/contributor-roles/supervision/">Supervision</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>Torres-Montúfar</surname>
            <given-names>Alejandro</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-5530-1319</uri>
          <xref ref-type="aff" rid="A4">4</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/formal-analysis/">Formal analysis</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>
          <role content-type="http://credit.niso.org/contributor-roles/software/">Software</role>
          <role content-type="http://credit.niso.org/contributor-roles/validation/">Validation</role>
          <role content-type="http://credit.niso.org/contributor-roles/visualization/">Visualization</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Universidad Científica del Sur, Ciencias de la Ingeniería, Carrera de Agronomía y Negocios, Campus Villa, Lima, Perú</addr-line>
        <institution>Herbario FES-Cuautitlán, Departamento de Ciencias Biológicas, Facultad de Estudios Superiores Cuautitlán, Universidad Nacional Autónoma de México</institution>
        <addr-line content-type="city">Cuautitlán Izcalli</addr-line>
        <country>Mexico</country>
        <uri content-type="ror">https://ror.org/01tmp8f25</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Botanischer Garten und Botanisches Museum Berlin, Freie Universität Berlin, Berlin, Germany</addr-line>
        <institution>University of Florida, School of Natural Resources and Environment</institution>
        <addr-line content-type="city">Gainesville</addr-line>
        <country>United States of America</country>
        <uri content-type="ror">https://ror.org/02y3ad647</uri>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">University of Florida, School of Natural Resources and Environment (SNRE), Gainesville, Florida, USA</addr-line>
        <institution>Botanischer Garten und Botanisches Museum Berlin, Freie Universität Berlin</institution>
        <addr-line content-type="city">Berlin</addr-line>
        <country>Germany</country>
        <uri content-type="ror">https://ror.org/046ak2485</uri>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Herbario FES-Cuautitlán, Departamento de Ciencias Biológicas, Facultad de Estudios Superiores Cuautitlán, Universidad Nacional Autónoma de México (FESC-UNAM), Cuautitlán Izcalli, México</addr-line>
        <institution>Universidad Científica del Sur, Ciencias de la Ingeniería, Carrera de Agronomía y Negocios</institution>
        <addr-line content-type="city">Lima</addr-line>
        <country>Peru</country>
        <uri content-type="ror">https://ror.org/04xr5we72</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Daniel B. Montesinos-Tubée (<email xlink:type="simple">dmontesinos@cientifica.edu.pe</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p><bold>Academic editor</bold>: André Simões</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>23</day>
        <month>04</month>
        <year>2026</year>
      </pub-date>
      <volume>159</volume>
      <issue>2</issue>
      <fpage>244</fpage>
      <lpage>254</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/5FE13F95-B3C1-5D9F-94BC-66729C1BBD25">5FE13F95-B3C1-5D9F-94BC-66729C1BBD25</uri>
      <history>
        <date date-type="received">
          <day>22</day>
          <month>10</month>
          <year>2025</year>
        </date>
        <date date-type="accepted">
          <day>15</day>
          <month>01</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Daniel B. Montesinos-Tubée, Cameron E.L. Dasher, Thomas Borsch, Alejandro Torres-Montúfar</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> – The newly described species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name>) from the Amazonas department of northern Peru represents an isolated lineage within the monophyletic genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>. This study aims to describe its distinctive morphological features and ecological context, assess its phylogenetic position, and discuss its implications for the diversification of the genus in the Andes.</p>
        <p><bold>Material and methods</bold> – Morphological observations were conducted on herbarium specimens and field collections from the type locality. Phylogenetic analyses were performed using nuclear and plastid DNA sequences from the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name>, with particular emphasis on <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>.</p>
        <p><bold>Key results</bold> – <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is morphologically distinct from its congeners by its acuminate leaf and sepal apices, hirsute-lanuginose seeds, and the presence of protective hooks surrounding the ovary. It inhabits humid, high-elevation grasslands known as “Jalca,” where it coexists with a diverse assemblage of montane species. The species appears to be endemic to the region and is currently known from only three populations. Its restricted distribution, combined with potential threats such as overgrazing and fire, supports a preliminary conservation assessment as Critically Endangered (<abbrev xlink:title="Critically Endangered">CR</abbrev>).</p>
        <p><bold>Conclusion</bold> – The discovery of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. highlights the underestimated diversity of Andean <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name> and provides evidence that Andean species form a grade within the diversification of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>. This pattern suggests a historical northward expansion of the lineage into the high mountains of Panama and Costa Rica.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>Amazonian</kwd>
        <kwd>endemism</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>taxonomy</kwd>
        <kwd>tropical Andes</kwd>
      </kwd-group>
      <funding-group>
        <award-group>
          <funding-source>
            <named-content content-type="funder_name">Alexander von Humboldt-Stiftung</named-content>
            <named-content content-type="funder_identifier">100005156</named-content>
            <named-content content-type="funder_ror">https://ror.org/012kf4317</named-content>
            <named-content content-type="funder_doi">http://doi.org/10.13039/100005156</named-content>
          </funding-source>
        </award-group>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="sec1">
      <title>Introduction</title>
      <p>The tropical Andes is recognized as one of the world’s richest regions in terms of biodiversity, and studies predict that many species have yet to be described there (<xref ref-type="bibr" rid="B44">Swenson et al. 2012</xref>; <xref ref-type="bibr" rid="B32">Ondo et al. 2024</xref>). This hotspot is believed to harbour approximately 45,000 species of vascular plants (<xref ref-type="bibr" rid="B31">Olson and Dinerstein 2002</xref>; <xref ref-type="bibr" rid="B22">Mittermeier et al. 2005</xref>; <xref ref-type="bibr" rid="B33">Orme et al. 2005</xref>), making it a critical area for ecological research and conservation. Despite its exceptional species richness, many aspects of its flora and ecosystems are still poorly understood, highlighting the need for further exploration and documentation. Such efforts are essential not only to enhance scientific knowledge but also to support strategies aimed at preserving this globally significant centre of biodiversity in the face of ongoing environmental changes and anthropogenic pressures. The <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name> family, commonly known as the madder or coffee family, is one of the largest families of flowering plants with a cosmopolitan distribution but a significant concentration in tropical regions (<xref ref-type="bibr" rid="B40">Robbrecht 1988</xref>; <xref ref-type="bibr" rid="B41">Robbrecht and Manen 2006</xref>; <xref ref-type="bibr" rid="B6">Bremer and Eriksson 2009</xref>) with 14,181–14,266 species in approximately 586–615 genera (<xref ref-type="bibr" rid="B37">Razafimandimbison and Rydin 2024a</xref>, <xref ref-type="bibr" rid="B39">2024b</xref>; <xref ref-type="bibr" rid="B49">Verstraete et al. 2025</xref>). In South America, the family is notably diverse and represents a significant component of South America’s plant biodiversity, especially in humid tropical regions like the Amazon. The tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name> Bercht. &amp; J.Presl is a pantropical lineage of 1411 species (<xref ref-type="bibr" rid="B49">Verstraete et al. 2025</xref>) and 86 genera (<xref ref-type="bibr" rid="B39">Razafimandimbison and Rydin 2024b</xref>), dominated by herbaceous and suffruticose taxa that frequently colonize disturbed environments (<xref ref-type="bibr" rid="B8">Delprete and Jardim 2012</xref>). In America, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name> are currently represented by 23 genera, including recently described genera such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Paganuccia">Paganuccia</tp:taxon-name-part></tp:taxon-name></italic> R.M.Salas (<xref ref-type="bibr" rid="B29">Nuñez-Florentin et al. 2022</xref>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Januaria">Januaria</tp:taxon-name-part></tp:taxon-name></italic> R.M.Salas &amp; Nuñez Florentin (<xref ref-type="bibr" rid="B30">Nuñez-Florentin et al. 2023</xref>).</p>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name> is one of the largest lineages of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name> with major Neotropical representatives from the Amazon lowlands (e.g. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Borreria">Borreria</tp:taxon-name-part></tp:taxon-name></italic> G.Mey, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hexasepalum">Hexasepalum</tp:taxon-name-part></tp:taxon-name></italic> Bartl. ex DC., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Diodia">Diodia</tp:taxon-name-part></tp:taxon-name></italic> Gronov., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Mitracarpus">Mitracarpus</tp:taxon-name-part></tp:taxon-name></italic> Zucc., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Galianthe">Galianthe</tp:taxon-name-part></tp:taxon-name></italic> Griseb., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Psyllocarpus">Psyllocarpus</tp:taxon-name-part></tp:taxon-name></italic> Mart. &amp; Zucc., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Spermacoce">Spermacoce</tp:taxon-name-part></tp:taxon-name></italic> L.) and a conspicuous Andean element such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> Schult. &amp; Schult.f. which is a small but ecologically important group inhabiting high-altitude regions of the Andes including the norward extension of the Cordillera de Mérida in Venezuela and the Talamanca mountain range in western Panama and Costa Rica (<xref ref-type="bibr" rid="B19">Mena 1990</xref>). Members of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> are cushion or mat-forming shrubs and constitute important components in montane and paramo ecosystems, adapted to cold and humid conditions (<xref ref-type="bibr" rid="B43">Sturm and Rangel 1985</xref>; <xref ref-type="bibr" rid="B20">Mena and Balslev 1986</xref>; <xref ref-type="bibr" rid="B19">Mena 1990</xref>; <xref ref-type="bibr" rid="B3">Andersson et al. 2002</xref>).</p>
      <p><xref ref-type="bibr" rid="B19">Mena (1990)</xref> recognized 15 species in his revision, and <xref ref-type="bibr" rid="B45">Taylor et al. (1993)</xref> listed eight species for Peru including <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="weberbaueri">weberbaueri</tp:taxon-name-part></tp:taxon-name></italic> K.Krause, that was not considered by <xref ref-type="bibr" rid="B19">Mena (1990)</xref>. <xref ref-type="bibr" rid="B46">Terrell (1999)</xref> merged the Mexican <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hedyotis">Hedyotis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="serpyllacea">serpyllacea</tp:taxon-name-part></tp:taxon-name></italic> Schltdl. into <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>, what was rejected by <xref ref-type="bibr" rid="B3">Andersson et al. (2002)</xref> and <xref ref-type="bibr" rid="B10">Groeninckx et al. (2009)</xref> who found this species as sister to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Bouvardia">Bouvardia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="ternifolia">ternifolia</tp:taxon-name-part></tp:taxon-name></italic> (Cav.) Schltdl., in a clade close to the traditional narrowly circumscribed <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name>, but not related to the assemblage of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hedyotis">Hedyotis</tp:taxon-name-part></tp:taxon-name></italic> L., and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Houstonia">Houstonia</tp:taxon-name-part></tp:taxon-name></italic> Gronov. as a more distant group in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name> sensu lato. Another species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="fasciculatum">fasciculatum</tp:taxon-name-part></tp:taxon-name></italic> (A.Gray) Terrell &amp; H.Rob. with a distribution in the northern deserts of Mexico and reaching into Texas was transferred from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Houstonia">Houstonia</tp:taxon-name-part></tp:taxon-name></italic> to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> by <xref ref-type="bibr" rid="B47">Terrell and Robinson (2011)</xref>.</p>
      <p>This group has drawn attention for its taxonomic diversity and endemism, particularly in Peru, Colombia, and Ecuador (<xref ref-type="bibr" rid="B3">Andersson et al. 2002</xref>). Morphologically, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> species are characterized by small, opposite leaves, interpetiolar stipules, and diminutive, often inconspicuous flowers (<xref ref-type="bibr" rid="B53">Wolff and Liede-Schumann 2007</xref>). Despite these similarities, the genus displays notable variations in floral disposition (pedicellate or sessile), leaf shape, margin pubescence, and seed hilum position (<xref ref-type="bibr" rid="B19">Mena 1990</xref>; <xref ref-type="bibr" rid="B47">Terrell and Robinson 2011</xref>), which have been key in distinguishing species within the genus.</p>
      <p>As part of the ongoing comprehensive investigation of the flora of the mountainous regions of northern Peru several collections of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> were made that at first glance showed a similar habit as other cushion or mat forming species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arenaria">Arenaria</tp:taxon-name-part></tp:taxon-name></italic> L. or <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Stellaria">Stellaria</tp:taxon-name-part></tp:taxon-name></italic> L. in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Caryophyllaceae">Caryophyllaceae</tp:taxon-name-part></tp:taxon-name>, which are frequent in the same habitats (e.g. <xref ref-type="bibr" rid="B13">Iamonico and Montesinos-Tubée 2023</xref>; <xref ref-type="bibr" rid="B23">Montesinos-Tubée and Iamonico 2023</xref>; <xref ref-type="bibr" rid="B24">Montesinos-Tubée and Borsch 2023</xref>). When analysing these specimens closely, they became evident as members of the coffee family, and subsequent phylogenetic analysis indicated that they correspond to a hitherto undescribed species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>. This paper aims to describe this new species as new for science, including a detailed morphological description, illustrations, and an evaluation of its phylogenetic position.</p>
    </sec>
    <sec sec-type="materials|methods" id="sec2">
      <title>Material and methods</title>
      <sec sec-type="Plant material" id="sec3">
        <title>Plant material</title>
        <p>The first author collected specimens in the highlands of the Central Andes from 2015 to 2025. Specimens were studied using appropriate literature (<xref ref-type="bibr" rid="B47">Terrell and Robinson 2011</xref>) and compared to material deposited at AMAZ, B, CPUN, HUT, MEXU, and USM (herbarium acronyms according to <xref ref-type="bibr" rid="B48">Thiers 2025</xref>). Additionally, images of type specimens from diverse species were checked via JSTOR Global Plants (<xref ref-type="bibr" rid="B16">JSTOR 2025</xref>). Since these collections did not match any of the known species in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>, they were hypothesized to represent an undescribed species.</p>
      </sec>
      <sec sec-type="Morphological studies" id="sec4">
        <title>Morphological studies</title>
        <p>The description presented here is based on our field observations, herbarium specimens, and spirit material, while the terminology follows <xref ref-type="bibr" rid="B14">Harris and Harris (2001)</xref>.The primary habit characteristics of collected specimens were recorded directly during fieldwork. Additional morphological features were examined using herbarium samples using Olympus SZX10 and NSZ-405 stereo microscopes (1X-4.5X magnification). To further document the specimens, detailed field photographs were also captured during the collection process.</p>
      </sec>
      <sec sec-type="Distribution and conservation status" id="sec5">
        <title>Distribution and conservation status</title>
        <p>The distribution maps were drawn using locality data from our collections using QGIS Desktop v.3.4.11 (<xref ref-type="bibr" rid="B36">QGIS Development Team 2021</xref>). The maps were prepared using cartography shapefiles acquired from <xref ref-type="bibr" rid="B18">MapBiomas Peru (2025)</xref>. Conservation status was assessed by applying the IUCN Red List Category criteria (<xref ref-type="bibr" rid="B15">IUCN 2024</xref>). We used the GeoCAT program (<xref ref-type="bibr" rid="B4">Bachman et al. 2011</xref>) to estimate the extent of occurrence (<abbrev xlink:title="extent of occurrence">EOO</abbrev>) and area of occupancy (<abbrev xlink:title="area of occupancy">AOO</abbrev>), based on 2 × 2 km cells.</p>
      </sec>
      <sec sec-type="Molecular laboratory works" id="sec6">
        <title>Molecular laboratory works</title>
        <p>Genomic DNA was extracted from silica-gel-dried or herbarium samples using the CTAB method (<xref ref-type="bibr" rid="B5">Borsch et al. 2003</xref>). The ITS region was amplified using the primers 18S 1830 and 26S 25 (<xref ref-type="bibr" rid="B28">Nickrent et al. 1994</xref>). PCR conditions were: 40 cycles of denaturation (60 s at 97°C), annealing (50 s at 53°C), extension (20 s at 72°C), and a final extension step (7 min at 72°C). The <italic>rps</italic>16 region was amplified using the primers rpsF and rpsR2 (<xref ref-type="bibr" rid="B34">Oxelman et al. 1997</xref>). PCR conditions were 90 s at 96°C, 1 min at 50°C, 90 s at 72°C, 35 cycles of 30 seconds at 96°C, 1 min at 50°C, 90 s at 72°C, and a final extension time of 10 min at 72°C. Sequencing was outsourced to Macrogen Europe using the BigDye® Terminator Kit on an ABI 3730xl Analyzer.</p>
      </sec>
      <sec sec-type="Sequence alignment and phylogenetic analysis" id="sec7">
        <title>Sequence alignment and phylogenetic analysis</title>
        <p>All sequences were manually aligned following the criteria proposed by <xref ref-type="bibr" rid="B17">Löhne and Borsch (2005)</xref>. The newly generated pherograms were edited and assembled using PhyDE v.0.995 (<xref ref-type="bibr" rid="B25">Müller et al. 2005</xref>). The matrix comprises 82 terminals, corresponding to 19 genera and 68 species of the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe" reg="Spermacoceae">Spermacoceae</tp:taxon-name-part></tp:taxon-name>. Of all the accessions, two correspond to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. generated for this study, the other 80 sequences were generated for previous studies and downloaded from GenBank (<xref ref-type="bibr" rid="B2">Andersson and Rova 1999</xref>; <xref ref-type="bibr" rid="B3">Andersson et al. 2002</xref>; <xref ref-type="bibr" rid="B53">Wolff and Liede-Schumann 2007</xref>; <xref ref-type="bibr" rid="B26">Neupane et al. 2015</xref>, <xref ref-type="bibr" rid="B27">2017</xref>; <xref ref-type="bibr" rid="B21">Miguel et al. 2018</xref>; <xref ref-type="bibr" rid="B9">Gibbons 2020</xref>). Taxon names with authors and GenBank accession numbers are presented in Suppl. material 1.</p>
        <p>For Bayesian inference (<abbrev xlink:title="Bayesian inference">BI</abbrev>), best-fit models of sequence evolution were selected for the molecular marker following the results of the Akaike Information Criterion in Modeltest v.3.6 (<xref ref-type="bibr" rid="B35">Posada and Crandall 1998</xref>) with parameters of 203 substitution schemes, +F base frequencies, +I and +G rate variation, nCat = 4, and SPR tree search. <abbrev xlink:title="Bayesian inference">BI</abbrev> analyses were conducted using MrBayes v.3.1.2 (<xref ref-type="bibr" rid="B12">Huelsenbeck and Ronquist 2001</xref>). Four independent Markov Chain Monte Carlo (<abbrev xlink:title="Markov Chain Monte Carlo">MCMC</abbrev>) runs were carried out, each with four parallel chains. Each chain was performed for 1,000,000 generations, saving one random tree every 1000 generations, the first 25% of the trees were discarded as burn-in. Trees were selected based on a 50% majority-rule consensus to estimate the posterior probabilities. A posterior probability (<abbrev xlink:title="posterior probability">PP</abbrev>) equal or greater than 0.90 was considered as well supported (<xref ref-type="bibr" rid="B1">Alfaro and Holder 2006</xref>). Visualization and node confidence were annotated in TreeGraph v.2.15.0 (<xref ref-type="bibr" rid="B42">Stöver and Müller 2010</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="Results and discussion" id="sec8">
      <title>Results and discussion</title>
      <sec sec-type="Comparative morphological studies" id="sec9">
        <title>Comparative morphological studies</title>
        <p>Morphological features place <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. unambiguously within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>. The mat-forming growth habit with slender and short herbaceous stems, together with the overall leaf and stipule morphology, conforms to the diagnostic features of the genus. Among the published species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is morphologically most similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic> (Ruiz &amp; Pav.) Standl., particularly in its mat-forming habit and general leaf form, which may cause initial confusion between both species. However, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. can be readily distinguished by a suite of diagnostic characters: it has acuminate leaf and sepal apices (vs acute or aristate in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>), lacks the long digitiform projections characteristic of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>, and exhibits longer stipules (1.3–2.0 vs 0.3–1.0 mm). Seed morphology provides an additional clear distinction, as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. has hirsute-lanuginose seeds, whereas <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic> presents finely scaly seeds. The new species also resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic> (Wedd.) Standl. in its growth habit and stipule morphology but differs by its purplish flowers and the hirsute-lanuginose texture of its seeds, compared to the glabrate seeds of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic>. A noteworthy character of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is the presence of two hooks protecting the ovary, a feature unique within the genus, which may represent an ecological adaptation to the harsh environmental conditions of the northern Jalca. These hooks could act as a physical barrier against water damage, reducing the risk of seed rotting under prolonged moisture, and provide mechanical reinforcement against raindrop impact and herbivory. The main morphological differences between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. and its most similar congeners are summarized in Table <xref ref-type="table" rid="T1">1</xref>.</p>
        <table-wrap id="T1" position="float" orientation="portrait">
          <label>Table 1.</label>
          <caption>
            <p>Morphological comparison of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. and related species.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Character</bold>
                </td>
                <td rowspan="1" colspan="1"><bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov</bold>.</td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
                <td rowspan="1" colspan="1">
                  <bold>
                    <italic>
                      <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name>
                    </italic>
                  </bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Leaf shape</bold>
                </td>
                <td rowspan="1" colspan="1">Ovate-lanceolate</td>
                <td rowspan="1" colspan="1">Linear</td>
                <td rowspan="1" colspan="1">Lanceolate to linear-lanceolate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Leaf apex</bold>
                </td>
                <td rowspan="1" colspan="1">Acuminate</td>
                <td rowspan="1" colspan="1">Acute or aristate</td>
                <td rowspan="1" colspan="1">Acute to rounded</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Sepal apex</bold>
                </td>
                <td rowspan="1" colspan="1">Acuminate</td>
                <td rowspan="1" colspan="1">Acute or aristate</td>
                <td rowspan="1" colspan="1">Oblong</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Stipule length</bold>
                </td>
                <td rowspan="1" colspan="1">1.3–2.0 mm</td>
                <td rowspan="1" colspan="1">0.3–1.0 mm</td>
                <td rowspan="1" colspan="1">1 mm</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Corolla tube length</bold>
                </td>
                <td rowspan="1" colspan="1">1.2–1.5 mm</td>
                <td rowspan="1" colspan="1">1 mm</td>
                <td rowspan="1" colspan="1">3.0–4.5 mm</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Flower colour</bold>
                </td>
                <td rowspan="1" colspan="1">White</td>
                <td rowspan="1" colspan="1">White</td>
                <td rowspan="1" colspan="1">White with purplish tones</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Seed ornamentation</bold>
                </td>
                <td rowspan="1" colspan="1">Hirsute-lanuginose</td>
                <td rowspan="1" colspan="1">Finely scaly</td>
                <td rowspan="1" colspan="1">Glabrate</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Ovary protective structures</bold>
                </td>
                <td rowspan="1" colspan="1">Two hooks protecting the ovary</td>
                <td rowspan="1" colspan="1">Absent</td>
                <td rowspan="1" colspan="1">Absent</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="Phylogenetic analysis" id="sec10">
        <title>Phylogenetic analysis</title>
        <p>The aligned matrix with all regions consisted of 1493 characters. The selected substitution model was GTR+I+G. Detailed matrix statistics regarding each marker contribution are given in Table <xref ref-type="table" rid="T2">2</xref>.</p>
        <table-wrap id="T2" position="float" orientation="portrait">
          <label>Table 2.</label>
          <caption>
            <p>Contribution of the different genomic regions to the total matrix.</p>
          </caption>
          <table>
            <tbody>
              <tr>
                <td rowspan="1" colspan="1"/>
                <td rowspan="1" colspan="1">
                  <bold>ITS</bold>
                </td>
                <td rowspan="1" colspan="1">
                  <bold><italic>rps</italic>16</bold>
                </td>
                <td rowspan="1" colspan="1">
                  <bold>Total</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Number of sequences</bold>
                </td>
                <td rowspan="1" colspan="1">71</td>
                <td rowspan="1" colspan="1">61</td>
                <td rowspan="1" colspan="1">82</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Aligned positions</bold>
                </td>
                <td rowspan="1" colspan="1">610</td>
                <td rowspan="1" colspan="1">883</td>
                <td rowspan="1" colspan="1">1493</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Potentially informative characters for parsimony</bold>
                </td>
                <td rowspan="1" colspan="1">218 (35.7%)</td>
                <td rowspan="1" colspan="1">206 (23.3%)</td>
                <td rowspan="1" colspan="1">424 (28.3%)</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">
                  <bold>Model selected for <abbrev xlink:title="Bayesian inference">BI</abbrev></bold>
                </td>
                <td rowspan="1" colspan="1">GTR+I+G</td>
                <td rowspan="1" colspan="1">GTR+I+G</td>
                <td rowspan="1" colspan="1"/>
              </tr>
            </tbody>
          </table>
        </table-wrap>
        <p>Our phylogenetic analysis recovered the new species as a member of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> with strong support (Fig. <xref ref-type="fig" rid="F1">1</xref>). Within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>, the new species is resolved as an independent branch separated from its closest morphological relatives, reinforcing its recognition as a separate species. However, the limited terminal sampling in our dataset prevents us from drawing definitive conclusions about its closest phylogenetic allies or the biogeographical history of the group. Despite these limitations, the congruence between the molecular results and the diagnostic morphological characters highlighted above underscores the importance of combining phylogenetic evidence with detailed comparative morphology for the delimitation of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> species from the Andean highlands.</p>
        <fig id="F1">
          <object-id content-type="doi">10.5091/plecevo.175775.figure1</object-id>
          <object-id content-type="arpha">B22D89EE-3F57-593D-8C47-485841609996</object-id>
          <label>Figure 1.</label>
          <caption>
            <p>Reconstructed Bayesian phylogenetic tree of the concatenated markers ITS and <italic>rps16</italic> showing the <abbrev xlink:title="posterior probability">PP</abbrev> values at the nodes. The samples of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. are marked in red. The scale bar represents the mean number of nucleotide substitutions per site.</p>
          </caption>
          <graphic xlink:href="plecevo-159-244-g001.jpg" id="oo_1600430.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1600430</uri>
          </graphic>
        </fig>
      </sec>
    </sec>
    <sec sec-type="Taxonomic treatment" id="sec11">
      <title>Taxonomic treatment</title>
      <tp:taxon-treatment>
        <tp:treatment-meta>
          <kwd-group>
            <label>Taxon classification</label>
            <kwd>
              <named-content content-type="kingdom">Plantae</named-content>
            </kwd>
            <kwd>
              <named-content content-type="order">Gentianales</named-content>
            </kwd>
            <kwd>
              <named-content content-type="family">Rubiaceae</named-content>
            </kwd>
          </kwd-group>
        </tp:treatment-meta>
        <tp:nomenclature>
          <tp:taxon-name><object-id content-type="arpha">52B067FA-F6A4-5F94-AF6D-EAF0F33CA3FF</object-id>
                		<tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part>
                	
                		<object-id content-type="ipni" xlink:type="simple">urn:lsid:ipni.org:names:77378619-1</object-id>
                	</tp:taxon-name>
          <tp:taxon-authority>Montesin., Borsch &amp; Torr.-Montúfar</tp:taxon-authority>
          <tp:taxon-status>sp. nov.</tp:taxon-status>
          <xref ref-type="fig" rid="F2">Figs 2</xref>
          <xref ref-type="fig" rid="F3">, 3</xref>
          <xref ref-type="fig" rid="F4">, 4</xref>
        </tp:nomenclature>
        <tp:treatment-sec sec-type="Type">
          <title>Type</title>
          <p>PERU – <bold>Amazonas department</bold> • Chachapoyas, Leymebamba, Laguna de los Cóndores track, antes de Abra el Rayo, scrubland vegetation, very humid; <named-content content-type="dwc:verbatimCoordinates">6°46’06”S, 77°46’37”W</named-content>; 2990 m; 21 Sep. 2023; <italic>D.B. Montesinos, C.E.L. Dasher &amp; E. Chávez 9695</italic>; holotype: HUT [HUT-64624]; isotype: B [B 10 1330351].</p>
          <fig id="F2">
            <object-id content-type="doi">10.5091/plecevo.175775.figure2</object-id>
            <object-id content-type="arpha">83DF0D20-439C-536D-941E-17E4F84E2D82</object-id>
            <label>Figure 2.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. <bold>A</bold>. Habit. <bold>B</bold>. Sterile stem. <bold>C</bold>. Stipule. <bold>D</bold>. Flower. <bold>E</bold>. Fruit. Drawn by Aldo Domínguez de la Torre based on <italic>Montesinos 9695</italic>.</p>
            </caption>
            <graphic xlink:href="plecevo-159-244-g002.jpg" id="oo_1600431.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1600431</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Diagnosis">
          <title>Diagnosis</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. can be recognized within the genus by the presence of two hooks protecting the ovary, a unique characteristic for this taxon and by the combination of acuminate leaf apex and hirsute-lanuginose seeds; it differs from the also herbaceous and mat-forming <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic> in leaf shape (ovate-lanceolate vs linear in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>) and sepal apex shape (acuminate vs acute or aristate in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>), the absence of long digitiform projections (present in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>), stipule length (1.3–2.0 vs 0.3–1.0 mm in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>), and seed ornamentation (hirsute-lanuginose vs finely scaly in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="filiforme">filiforme</tp:taxon-name-part></tp:taxon-name></italic>). It also differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic>, another mat forming species by its sepal apex shape (oblong in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic>), corolla tube length (1.2–1.5 vs 3.0–4.5 mm long in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic>), and seed ornamentation (hirsute-lanuginose vs glabrous seeds in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="muticum">muticum</tp:taxon-name-part></tp:taxon-name></italic>).</p>
          <fig id="F3">
            <object-id content-type="doi">10.5091/plecevo.175775.figure3</object-id>
            <object-id content-type="arpha">5C4527B2-4911-5433-A7B2-183E88209710</object-id>
            <label>Figure 3.</label>
            <caption>
              <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. <bold>A</bold>. Habit. <bold>B</bold>. Flowers. <bold>C</bold>. Fruits (black arrows). Photographs by Daniel Montesinos.</p>
            </caption>
            <graphic xlink:href="plecevo-159-244-g003.jpg" id="oo_1600432.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1600432</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Description">
          <title>Description</title>
          <p>Perennial <underline>herb</underline> with a decumbent, prostrate, or erect growth habit, forming a dense and ramified mat measuring approximately 5–10 cm in diameter. <underline>Roots</underline> subwoody-fibrous. <underline>Stems</underline> ranging from 5 to 8 cm in length, display frequent branching and are often concealed by older leaves; bark dark red hue, young shoots exhibit shades of dark green to pale red. <underline>Internodes</underline> short (2.0–3.5 mm) and narrow (0.2–0.3 mm), scarcely covered by irregular brown protuberances less than 0.05 mm in length, striate, deeply sulcate with 9–11 ribs. <underline>Stipules</underline> oblong-lanceolate, translucid, 1.3–2 × 0.8–1.8 mm, acuminate apex, irregular dentate margins, base truncate. <underline>Leaves</underline> opposite, ovate-lanceolate, 1.9–2.7 × 0.5–0.7 mm, sessile, apically subrevolute, slightly bulging midnerve, lamina dark green, thick, densely covered by sericeous trichomes on the underside and margins, sparsely and glandulous on the upperside, margin trichomes are whitish, measuring 0.10–0.25 mm long, multicellular, 18 to 40 per margin side, leaf apex largely acuminate, hirsute, hooked, with a long linear to sublinear ciliate hair, 0.5–0.8 mm long that tends to turn yellowish red with age, persisting on older foliage, leaf base truncate, narrow, densely hirsute on both sides and margins. <underline>Flowers</underline> solitary, axillary or terminal, tubular to shortly campanulate, tube 1.2–1.5 mm long, sessile or pedicellate, pedicels 0.1–0.2 mm long, sepals 4, ovate-lanceolate, 2.0–2.3 × 0.5–0.8 mm, covered with a thin translucid ovate stipule, densely hirsute on margins and upper surface, bearing a whitish acumen of approximately the same size as in the leaves, 2.0–2.2 mm long; petals 4, entire, white translucid, with a unique lobule, ovate-lanceolate, 1.8–2.0 × 0.8–1.0 mm, glabrous, hidden by the concave sepal, and fused with the stamens at the base. <underline>Androecium</underline> with 10 stamens, spreading and slightly curved filaments, 1.2–1.4 mm including the roundish and reddish anthers, which exceed the size of the ovary. <underline>Gynoecium</underline> 1 mm long, ovary ovate, 0.9–0.8 × 0.4–0.6 mm, with a slightly hirsute surface, ovary protected by two hooks. <underline>Fruits</underline> capsular, roundish, persistent, sparsely hirsute, otherwise glandulose, 1.1–1.3 mm in diameter, five carpels. <underline>Seeds</underline> about 6 to 8 per capsule, ovate to triangular, 0.4–0.6 mm in diameter, densely pubescent, trichomes dark maroon coloured and glandulose.</p>
          <fig id="F4">
            <object-id content-type="doi">10.5091/plecevo.175775.figure4</object-id>
            <object-id content-type="arpha">FAC4897A-E631-5729-89C2-CC9A87668AA4</object-id>
            <label>Figure 4.</label>
            <caption>
              <p>Distribution map of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
            </caption>
            <graphic xlink:href="plecevo-159-244-g004.jpg" id="oo_1600433.jpg">
              <uri content-type="original_file">https://binary.pensoft.net/fig/1600433</uri>
            </graphic>
          </fig>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Distribution">
          <title>Distribution</title>
          <p>This species is exclusively documented in the Amazonas department, Peru. This area is situated within the Amotape–Huancabamba Zone, a major biodiversity hotspot extending from the Pacific coast, across the Andean cordilleras, to the tropical lowlands of southern Ecuador and much of northern Peru (<xref ref-type="bibr" rid="B50">Weigend 2002</xref>, <xref ref-type="bibr" rid="B51">2004</xref>) with many narrowly endemic species concentrated (e.g. <xref ref-type="bibr" rid="B50">Weigend 2002</xref>; <xref ref-type="bibr" rid="B52">Weigend et al. 2005</xref>; <xref ref-type="bibr" rid="B11">Henning et al. 2019</xref>). It primarily thrives in the Jalca ecosystems, which are characterized by persistent humidity in forest patches amidst wet grasslands. These ecosystems are typically found at elevations ranging from approximately 2,990 to 3,791 m. Despite extensive surveys conducted by the first author across all Peruvian herbaria, no herbarium collections or additional specimens have been found, underscoring its status as an endemic species in northern Peru. The Jalca region, which borders dense montane tropical forests and transitions to lowland tropical forests, receives continuous rainfall year-round. Notably, the highest point of the collection site, around 3,800 m in Pampa del Rayo on the route to the Laguna de los Cóndores, receives an estimated 2,500 mm of precipitation annually.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Phenology">
          <title>Phenology</title>
          <p>The new entity exhibits a flexible phenological pattern finely tuned to the extreme rainfall conditions of the northern Jalca in Peru, an ecosystem characterized by persistent and intense precipitation. Flowering and fruiting periods are highly variable, primarily influenced by the amount and timing of rainfall. Typically, flowering begins as early as August, coinciding with the onset of consistent rains, and may extend into October, depending on the intensity and duration of the wet season. This variability suggests a strong ecological adaptation to the region’s erratic precipitation patterns, enabling the species to optimize reproductive success during favourable moisture conditions.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Vernacular name">
          <title>Vernacular name</title>
          <p>“Atunina”, name used by the locals in the communities of Atuén and Dos de Mayo of the Leymebamba district.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Etymology">
          <title>Etymology</title>
          <p>Named after the Leymebamba district, several populations were identified in the vicinity of Laguna de los Cóndores. Although the lagoon itself lies just within San Martín (province of Huallaga) the type specimen’s collection site is clearly located in Amazonas in the Amotape–Huancabamba Zone as defined by <xref ref-type="bibr" rid="B7">Casper et al. (2020)</xref>. This area represents a significant habitat for the species, emphasizing its ecological importance and potential for further research. The diverse environments found in the Jalca ecosystems in the uppermost west topographical areas of the lagoon may contribute to the species unique adaptations and distribution patterns.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Preliminary IUCN conservation assessment">
          <title>Preliminary IUCN conservation assessment</title>
          <p>The species is given a Red List status of Critically Endangered [<abbrev xlink:title="Critically Endangered">CR</abbrev> B1ab(iii)+2ab(iii)]. So far, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. has been identified in only three locations, making it a relatively uncommon species. The extent of occurrence (<abbrev xlink:title="extent of occurrence">EOO</abbrev>) is of 1.2 km<sup>2</sup> and the area of occupancy (<abbrev xlink:title="area of occupancy">AOO</abbrev>) is of 8 km<sup>2</sup>. Both <abbrev xlink:title="extent of occurrence">EOO</abbrev> and <abbrev xlink:title="area of occupancy">AOO</abbrev> fall within the limits of the Critically Endangered (<abbrev xlink:title="Critically Endangered">CR</abbrev>) category under subcriteria B1 and B2. According to the IUCN criteria and categories (<xref ref-type="bibr" rid="B15">IUCN 2024</xref>), it meets the conditions for the <abbrev xlink:title="Critically Endangered">CR</abbrev> category. The preferred habitats for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. are the summit slopes of Jalca ecosystems, adjacent to lowland tropical ecosystems in northern Peru. These mountain slopes face vulnerabilities from factors such as overgrazing, changes in annual rainfall, and fire, all of which could reduce the distribution area of this species. During a field visit by the first author in 2023, extensive burning of these ecosystems was observed. Despite their natural resistance to fire, rapid environmental changes threaten biodiversity in the region. Furthermore, recent fires in September 2024 have devastated much of the habitat where this species grows, leading to concerns about its potential disappearance unless immediate conservation protocols are implemented by the state.</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Additional material examined">
          <title>Additional material examined</title>
          <p>PERU – <bold>Amazonas Department</bold> • Chachapoyas Province, Leymebamba District, Camino a Laguna Yanacocha, humid grasslands, forest patches, continuously humid; 3791 m; 23 Sep. 2023; <italic>D.B. Montesinos, C.E.L. Dasher &amp; E. Chávez 9706</italic>; HUT [HUT-64623], B [B 10 1330350] • Jalca ecosystem, highly moist, dense scrubland with tussock grasses; 3760 m; 22 Jul. 2024; <italic>D.B. Montesinos, C.E.L. Dasher &amp; E. Chávez 10400</italic>; AMAZ, B [B 10 1330352].</p>
        </tp:treatment-sec>
        <tp:treatment-sec sec-type="Phylogenetic position">
          <title>Phylogenetic position</title>
          <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> was retrieved as monophyletic and highly supported either in combined analysis (<abbrev xlink:title="posterior probability">PP</abbrev> = 1.00) and in each individual molecular marker analysis (results not shown). Both samples of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. form a clade with high Bayesian posterior probabilities in the combined analysis (<abbrev xlink:title="posterior probability">PP</abbrev> = 1.00) and in the ITS region (<abbrev xlink:title="posterior probability">PP</abbrev> = 1.00) (results not shown), only one sample amplified for the <italic>rps</italic>16 marker. Furthermore, the phylogeny also shows that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. is closest to a clade formed by several Andean species, however, the sampling of both characters and terminals prevents establishing a biogeographic hypothesis and this molecular analysis only confirms the existence of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">A.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leymebambense">leymebambense</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. as a unique lineage within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Arcytophyllum">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>.</p>
        </tp:treatment-sec>
      </tp:taxon-treatment>
    </sec>
    <sec sec-type="Conclusions" id="sec12">
      <title>Conclusions</title>
      <p>The importance of this work lies in the identification and formal description of a new lineage from the Andean region, belonging to a genus of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Rubiaceae">Rubiaceae</tp:taxon-name-part></tp:taxon-name> characteristic of the high-elevation zones of South America. This contribution not only expands the known diversity of the group but also underscores the value of continued botanical exploration in understudied areas. Moreover, it reinforces the relevance of integrative systematics, in which phylogenetic analyses are combined with detailed morphological studies to achieve a deeper understanding of the evolutionary patterns and diversification processes shaping these montane ecosystems.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>This work was conducted as part of the postdoctoral project of the first author, sponsored by the Alexander von Humboldt-Stiftung through a Georg Forster Research Fellowship. Thanks are due to Anna Guengerich for her fieldwork support in the Amazonas region, financed by US National Science Foundation, award #2019425. We also thank the residents of Leymebamba, Dos de Mayo, and Atuén for their assistance in the field in the Amazonas region. Additionally, funding was previously obtained through J.S. Ingham. We express our gratitude to E.F. Rodríguez from the Herbario HUT (Universidad Nacional de Trujillo), and P. Hein and S. Bernhard (Botanischer Garten und Botanisches Museum Berlin), for their support with herbarium consultations. The independent illustrator Aldo Domínguez de la Torre made the illustration of the type. The various botanical collections carried out are covered by collection permits granted by the Peruvian State: Letter N° D000375-2023-MIDAGRI-SERFOR-DGGSPFFS-DGSPF, Letter N° RD-000130-2023-DGGSPFFS-DGSPF, and Letter N° RD-000026-2024-DGGSPFFS-DGSPF.</p>
    </ack>
    <ref-list>
      <title>References</title>
      <ref id="B1">
        <mixed-citation>Alfaro ME, Holder MT (2006) The posterior and the prior in Bayesian phylogenetics. Annual Review of Ecology, Evolution, and Systematics 37: 19–42. <ext-link xlink:href="10.1146/annurev.ecolsys.37.091305.110021" ext-link-type="doi">https://doi.org/10.1146/annurev.ecolsys.37.091305.110021</ext-link></mixed-citation>
      </ref>
      <ref id="B2">
        <mixed-citation>Andersson L, Rova JHE (1999) The <italic>rps</italic>16 intron and the phylogeny of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rubioideae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>). Plant Systematics and Evolution 214: 161–186. <ext-link xlink:href="10.1007/BF00985737" ext-link-type="doi">https://doi.org/10.1007/BF00985737</ext-link></mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation>Andersson L, Rova JHE, Guarin FA (2002) Relationships, circumscription, and biogeography of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>) based on evidence from cpDNA. Brittonia 54(1): 40–49. <ext-link xlink:href="10.1663/0007-196X(2002)054" ext-link-type="doi">https://doi.org/10.1663/0007-196X(2002)054</ext-link>[0040:RCABOA]2.0.CO;2</mixed-citation>
      </ref>
      <ref id="B4">
        <mixed-citation>Bachman S, Moat J, Hill AW, de la Torre J, Scott B (2011) Supporting red list threat assessments with GeoCAT: geospatial conservation assessment tool. ZooKeys 150: 111–126. <ext-link xlink:href="10.3897/zookeys.150.2109" ext-link-type="doi">https://doi.org/10.3897/zookeys.150.2109</ext-link></mixed-citation>
      </ref>
      <ref id="B5">
        <mixed-citation>Borsch T, Hilu KW, Quandt D, Wilde V, Neinhuis C, Barthlott W (2003) Noncoding plastid <italic>trnT–trnF</italic> sequences reveal a well resolved phylogeny of basal angiosperms. Journal of Evolutionary Biology 16(4): 558–576. <ext-link xlink:href="10.1046/j.1420-9101.2003.00577.x" ext-link-type="doi">https://doi.org/10.1046/j.1420-9101.2003.00577.x</ext-link></mixed-citation>
      </ref>
      <ref id="B6">
        <mixed-citation>Bremer B, Eriksson T (2009) Time tree of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: phylogeny and dating the family, subfamilies and tribes. International Journal of Plant Sciences 170: 766–793. <ext-link xlink:href="10.1086/599077" ext-link-type="doi">https://doi.org/10.1086/599077</ext-link></mixed-citation>
      </ref>
      <ref id="B7">
        <mixed-citation>Casper SJ, Bussmann RW, Henning T (2020) <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Pinguicula</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">rosmarieae</tp:taxon-name-part></tp:taxon-name></italic> Casper, Bussmann &amp; T.Henning (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Lentibulariaceae</tp:taxon-name-part></tp:taxon-name>), a new butterwort from the Amotape-Huancabamba Zone (northern Peru). PhytoKeys 140: 107–123. <ext-link xlink:href="10.3897/phytokeys.140.49529" ext-link-type="doi">https://doi.org/10.3897/phytokeys.140.49529</ext-link></mixed-citation>
      </ref>
      <ref id="B8">
        <mixed-citation>Delprete PG, Jardim JG (2012) Systematics, taxonomy and floristics of Brazilian <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: an overview about the current status and future challenges. Rodriguésia 63: 101–128. <ext-link xlink:href="10.1590/S2175-78602012000100009" ext-link-type="doi">https://doi.org/10.1590/S2175-78602012000100009</ext-link></mixed-citation>
      </ref>
      <ref id="B9">
        <mixed-citation>Gibbons KL (2020) <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Hedyotis</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Oldenlandia</tp:taxon-name-part></tp:taxon-name></italic> and related genera (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name>) in Australia: new genera and new combinations in an Asian-Australian-Pacific lineage. Taxon 69: 515–542. <ext-link xlink:href="10.1002/tax.12236" ext-link-type="doi">https://doi.org/10.1002/tax.12236</ext-link></mixed-citation>
      </ref>
      <ref id="B10">
        <mixed-citation>Groeninckx I, Dessein S, Ochoterena H, Persson C, Motley TJ, Kårehed J, Bremer B, Huysmans S, Smets E (2009) Phylogeny of the herbaceous tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>) based on plastid DNA data. Annals of the Missouri Botanical Garden 96: 109–132. <ext-link xlink:href="10.3417/2006201" ext-link-type="doi">https://doi.org/10.3417/2006201</ext-link></mixed-citation>
      </ref>
      <ref id="B11">
        <mixed-citation>Henning T, Acuña Castillo RH, Rodríguez Rodríguez EF, Garcia Llatas LF, Weigend M (2019) A new striking and critically endangered species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Nasa</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Loasaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Cornales</tp:taxon-name-part></tp:taxon-name>) from North Peru. PhytoKeys 121: 13–28. <ext-link xlink:href="10.3897/phytokeys.121.33927" ext-link-type="doi">https://doi.org/10.3897/phytokeys.121.33927</ext-link></mixed-citation>
      </ref>
      <ref id="B12">
        <mixed-citation>Huelsenbeck JP, Ronquist F (2001) MRBAYES: Bayesian inference of phylogenetic trees. Bioinformatics 17: 754–755. <ext-link xlink:href="10.1093/bioinformatics/17.8.754" ext-link-type="doi">https://doi.org/10.1093/bioinformatics/17.8.754</ext-link></mixed-citation>
      </ref>
      <ref id="B13">
        <mixed-citation>Iamonico D, Montesinos-Tubée DB (2023) The genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Paronychia</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Caryophyllaceae</tp:taxon-name-part></tp:taxon-name>) in South America: nomenclatural review and taxonomic notes with the description of a new species from North Peru. Plants 12: 1064. <ext-link xlink:href="10.3390/plants12051064" ext-link-type="doi">https://doi.org/10.3390/plants12051064</ext-link></mixed-citation>
      </ref>
      <ref id="B14">
        <mixed-citation>Harris JG, Harris MW (2001) Plant Identification Terminology: an Illustrated Glossary. Second edition. Spring Lake Publishing, Spring Lake.</mixed-citation>
      </ref>
      <ref id="B15">
        <mixed-citation>IUCN (2024) Guidelines for Using the IUCN Red List Categories and Criteria. Version 16. Prepared by the Standards and Petitions Committee. <ext-link xlink:href="https://www.iucnredlist.org/resources/redlistguidelines" ext-link-type="uri">https://www.iucnredlist.org/resources/redlistguidelines</ext-link> [accessed 18.11.2025]</mixed-citation>
      </ref>
      <ref id="B16">
        <mixed-citation>JSTOR (2025) JSTOR Global Plants Database. <ext-link xlink:href="https://plants.jstor.org" ext-link-type="uri">https://plants.jstor.org</ext-link> [accessed 10.09.2025]</mixed-citation>
      </ref>
      <ref id="B17">
        <mixed-citation>Löhne C, Borsch T (2005) Molecular evolution and phylogenetic utility of the <italic>petD</italic> group II intron: a case study in basal angiosperms. Molecular Biology and Evolution 22: 317–332. <ext-link xlink:href="10.1093/molbev/msi019" ext-link-type="doi">https://doi.org/10.1093/molbev/msi019</ext-link></mixed-citation>
      </ref>
      <ref id="B18">
        <mixed-citation>MapBiomas Peru (2025) Shapefiles. <ext-link xlink:href="https://peru.mapbiomas.org/capas/" ext-link-type="uri">https://peru.mapbiomas.org/capas/</ext-link> [accessed 16.11.2025]</mixed-citation>
      </ref>
      <ref id="B19">
        <mixed-citation>Mena P (1990) A revision of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Hedyotideae</tp:taxon-name-part></tp:taxon-name>). Memoirs of the New York Botanical Garden 60: 1–26.</mixed-citation>
      </ref>
      <ref id="B20">
        <mixed-citation>Mena P, Balslev H (1986) Comparación entre la vegetación de los Páramos y el cinturón afroalpino. Reports from the Botanical Institute, University of Aarhus 12: 1–54.</mixed-citation>
      </ref>
      <ref id="B21">
        <mixed-citation>Miguel LM, Sobrado SV, Janssens S, Dessein S, Cabral EL (2018) The monotypic Brazilian genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Diacrodon</tp:taxon-name-part></tp:taxon-name></italic> is a synonym of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Borreria</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>): morphological and molecular evidences. Anais da Academia Brasileira de Ciências 90(2): 1397–1415. <ext-link xlink:href="10.1590/0001-3765201820170314" ext-link-type="doi">https://doi.org/10.1590/0001-3765201820170314</ext-link></mixed-citation>
      </ref>
      <ref id="B22">
        <mixed-citation>Mittermeier RA, Gil PR, Hoffman M, Pilgrim J, Brooks T, Mittermeier CG, Lamoreux J, de Fonseca GAB (2005) Hotspots Revisited: Earth’s Biologically Richest and Most Endangered Terrestrial Ecoregions. University of Chicago Press, Chicago.</mixed-citation>
      </ref>
      <ref id="B23">
        <mixed-citation>Montesinos-Tubée DB, Iamonico D (2023) Typifications of Andean <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arenaria</tp:taxon-name-part></tp:taxon-name></italic> names (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Caryophyllaceae</tp:taxon-name-part></tp:taxon-name>), with a nomen novum. Phytotaxa 579(1): 54–60. <ext-link xlink:href="10.11646/phytotaxa.579.1.6" ext-link-type="doi">https://doi.org/10.11646/phytotaxa.579.1.6</ext-link></mixed-citation>
      </ref>
      <ref id="B24">
        <mixed-citation>Montesinos-Tubée DB, Borsch T (2023) Molecular phylogenetics and morphology reveal the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Plettkea</tp:taxon-name-part></tp:taxon-name></italic> lineage including several members of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arenaria</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Pycnophyllopsis</tp:taxon-name-part></tp:taxon-name></italic> to be a clade of 21 South American species nested within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Stellaria</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Caryophyllaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Alsineae</tp:taxon-name-part></tp:taxon-name>). Willdenowia 53(3): 115–148. <ext-link xlink:href="10.3372/wi.53.53301" ext-link-type="doi">https://doi.org/10.3372/wi.53.53301</ext-link></mixed-citation>
      </ref>
      <ref id="B25">
        <mixed-citation>Müller K, Quandt D, Müller J, Neinhuis C (2005) PhyDE 0.995: Phylogenetic data editor. <ext-link xlink:href="http://www.phyde.de/" ext-link-type="uri">http://www.phyde.de/</ext-link> [accessed 19.01.2026]</mixed-citation>
      </ref>
      <ref id="B26">
        <mixed-citation>Neupane S, Dessein S, Wikström N, Lewis PO, Long C, Bremer B, Motley TJ (2015) The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Hedyotis</tp:taxon-name-part></tp:taxon-name>-<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Oldenlandia</tp:taxon-name-part></tp:taxon-name></italic> complex (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name>) in Asia and the Pacific: phylogeny revisited with new generic delimitations. Taxon 64: 299–322. <ext-link xlink:href="10.12705/642.8" ext-link-type="doi">https://doi.org/10.12705/642.8</ext-link></mixed-citation>
      </ref>
      <ref id="B27">
        <mixed-citation>Neupane S, Lewis PO, Dessein S, Shanks H, Paudyal S, Lens F (2017) Evolution of woody life form on tropical mountains in the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>). American Journal of Botany 104: 419–438. <ext-link xlink:href="10.3732/ajb.1600248" ext-link-type="doi">https://doi.org/10.3732/ajb.1600248</ext-link></mixed-citation>
      </ref>
      <ref id="B28">
        <mixed-citation>Nickrent DL, Schuette KP, Starr EM (1994) A molecular phylogeny of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arceuthobium</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Viscaceae</tp:taxon-name-part></tp:taxon-name>) based on nuclear ribosomal DNA internal transcribed spacer sequences. American Journal of Botany 81: 1149–1160. <ext-link xlink:href="10.1002/j.1537-2197.1994.tb15609.x" ext-link-type="doi">https://doi.org/10.1002/j.1537-2197.1994.tb15609.x</ext-link></mixed-citation>
      </ref>
      <ref id="B29">
        <mixed-citation>Nuñez-Florentin M, Salas RM, Carmo JAM, Cabral EL, Dessein S, Janssens SB (2022) <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Paganuccia</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">icatuensis</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>), a new genus and species from Bahia, Brazil, with a key to all the genera of the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name> in the Americas. Taxon 71(3): 630–649. <ext-link xlink:href="10.1002/tax.12651" ext-link-type="doi">https://doi.org/10.1002/tax.12651</ext-link></mixed-citation>
      </ref>
      <ref id="B30">
        <mixed-citation>Nuñez-Florentin M, Florentin JE, Salas RM, Dessein S, Janssens SB (2023) Multi-loci phylogeny and morphological evidence support the recognition of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Januaria</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Spermacoceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>), a new monospecific genus endemic to the North of Minas Gerais (Brazil). Anais da Academia Brasileira de Ciências 95(1): e20211601. <ext-link xlink:href="10.1590/0001-3765202320211601" ext-link-type="doi">https://doi.org/10.1590/0001-3765202320211601</ext-link></mixed-citation>
      </ref>
      <ref id="B31">
        <mixed-citation>Olson DM, Dinerstein E (2002) The Global 200: priority ecoregions for global conservation. Annals of the Missouri Botanical Garden 89: 199–224. <ext-link xlink:href="10.2307/3298564" ext-link-type="doi">https://doi.org/10.2307/3298564</ext-link></mixed-citation>
      </ref>
      <ref id="B32">
        <mixed-citation>Ondo I, Dhanjal-Adams KL, Pironon S, Silvestro D,Colli-Silva M, Deklerck V, Grace OM, Monro AK, Nicolson N, Walker B, Antonelli A (2024) Plant diversity darkspots for global collection priorities. New Phytologist 244: 719–733. <ext-link xlink:href="10.1111/nph.20024" ext-link-type="doi">https://doi.org/10.1111/nph.20024</ext-link></mixed-citation>
      </ref>
      <ref id="B33">
        <mixed-citation>Orme CDL, Davies RG, Burgess M, Eigenbrod F, Pickup N, Olson VA, Webster AJ, Ding T-S, Rasmussen PC, Ridgely RS, Stattersfield AJ, Bennett PM, Blackburn TM, Gaston KJ, Owens IPF (2005) Global hotspots of species richness are not congruent with endemism or threat. Nature 436: 1016–1019. <ext-link xlink:href="10.1038/nature03850" ext-link-type="doi">https://doi.org/10.1038/nature03850</ext-link></mixed-citation>
      </ref>
      <ref id="B34">
        <mixed-citation>Oxelman B, Lidén M, Berglund D (1997) Chloroplast <italic>rps</italic>16 intron phylogeny of the tribe Sileneae (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Caryophyllaceae</tp:taxon-name-part></tp:taxon-name>). Plant Systematics and Evolution 206(1): 393–410. <ext-link xlink:href="10.1007/BF00987959" ext-link-type="doi">https://doi.org/10.1007/BF00987959</ext-link></mixed-citation>
      </ref>
      <ref id="B35">
        <mixed-citation>Posada D, Crandall KA (1998) MODELTEST: testing the model of DNA substitution. Bioinformatics 14(9): 817–818. <ext-link xlink:href="10.1093/bioinformatics/14.9.817" ext-link-type="doi">https://doi.org/10.1093/bioinformatics/14.9.817</ext-link></mixed-citation>
      </ref>
      <ref id="B36">
        <mixed-citation>QGIS Development Team (2021) QGIS Geographic Information System. Version 3.4.11. Open Source Geospatial Foundation Project. <ext-link xlink:href="https://www.qgis.org" ext-link-type="uri">https://www.qgis.org</ext-link> [accessed 30.03.2026]</mixed-citation>
      </ref>
      <ref id="B37">
        <mixed-citation>Razafimandimbison SG, Rydin C (2024a) Phylogeny and classification of the coffee family (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Gentianales</tp:taxon-name-part></tp:taxon-name>): overview and outlook. Taxon 73: 673–717.</mixed-citation>
      </ref>
      <ref id="B38">
        <mixed-citation>
          <ext-link xlink:href="10.1002/tax.13167" ext-link-type="doi">https://doi.org/10.1002/tax.13167</ext-link>
        </mixed-citation>
      </ref>
      <ref id="B39">
        <mixed-citation>Razafimandimbison SG, Rydin C (2024b) Addendum and corrigendum to: Phylogeny and classification of the coffee family (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Gentianales</tp:taxon-name-part></tp:taxon-name>): overview and outlook [in Taxon 73: 673–717. 2024]. Taxon 73(6): 1556–1557. <ext-link xlink:href="10.1002/tax.13293" ext-link-type="doi">https://doi.org/10.1002/tax.13293</ext-link></mixed-citation>
      </ref>
      <ref id="B40">
        <mixed-citation>Robbrecht E (1988) Tropical woody <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>. Opera Botanica Belgica 1: 1–272.</mixed-citation>
      </ref>
      <ref id="B41">
        <mixed-citation>Robbrecht E, Manen JF (2006) The major evolutionary lineages of the coffee family (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>, angiosperms). Combined analysis (nDNA and cpDNA) to infer the position of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Coptosapelta</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Luculia</tp:taxon-name-part></tp:taxon-name></italic>, and supertree construction based on <italic>rbcL</italic>, <italic>rps16</italic>, <italic>trnL-trnF</italic> and <italic>atpB-rbcL</italic> data. A new classification in two subfamilies, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Cinchonoideae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Rubioideae</tp:taxon-name-part></tp:taxon-name>. Systematics and Geography of Plants 76: 85–146. <ext-link xlink:href="https://www.jstor.org/stable/20649700" ext-link-type="uri">https://www.jstor.org/stable/20649700</ext-link> [accessed 28.01.2026]</mixed-citation>
      </ref>
      <ref id="B42">
        <mixed-citation>Stöver BC, Müller KF (2010) TreeGraph 2: combining and visualizing evidence from different phylogenetic analyses. BMC Bioinformatics 11(7). <ext-link xlink:href="10.1186/1471-2105-11-7" ext-link-type="doi">https://doi.org/10.1186/1471-2105-11-7</ext-link></mixed-citation>
      </ref>
      <ref id="B43">
        <mixed-citation>Sturm H, Rangel O (1985) Ecología de los Páramos Andinos: una Visión Preliminar Integrada. Biblioteca J. Triana 9. Editorial Guadalupe, Bogotá.</mixed-citation>
      </ref>
      <ref id="B44">
        <mixed-citation>Swenson JJ, Young BE, Beck S, et al. (2012) Plant and animal endemism in the eastern Andean slope: challenges to conservation. BMC Ecology 12: 1. <ext-link xlink:href="10.1186/1472-6785-12-1" ext-link-type="doi">https://doi.org/10.1186/1472-6785-12-1</ext-link></mixed-citation>
      </ref>
      <ref id="B45">
        <mixed-citation>Taylor CM, Pool A, Kirkbride JH (1993) <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>. In: Brako L, Zarucchi J, Plumptre AJ (Eds) Catalogue of the Flowering Plants and Gymnosperms of Peru. Monographs in Systematic Botany from the Missouri Botanical Garden 45: 1001–1053.</mixed-citation>
      </ref>
      <ref id="B46">
        <mixed-citation>Terrell EE (1999) Morphology and taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arcytophyllum</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">serpyllaceum</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>), a transfer from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Hedyotis</tp:taxon-name-part></tp:taxon-name></italic>. Novon 9(2): 263–264. <ext-link xlink:href="10.2307/3391813" ext-link-type="doi">https://doi.org/10.2307/3391813</ext-link></mixed-citation>
      </ref>
      <ref id="B47">
        <mixed-citation>Terrell EE, Robinson H (2011) Fasciculate morphology, seed variation, and distributional notes in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>). Journal of the Botanical Research Institute of Texas 5(2): 587–594. <ext-link xlink:href="https://www.jstor.org/stable/41972308" ext-link-type="uri">https://www.jstor.org/stable/41972308</ext-link> [accessed 28.01.2026]</mixed-citation>
      </ref>
      <ref id="B48">
        <mixed-citation>Thiers B (2025) Index Herbariorum: a global directory of public herbaria and associated staff. New York Botanical Garden’s Virtual Herbarium. <ext-link xlink:href="https://sweetgum.nybg.org/science/ih/" ext-link-type="uri">https://sweetgum.nybg.org/science/ih/</ext-link> [accessed 13.11.2025]</mixed-citation>
      </ref>
      <ref id="B49">
        <mixed-citation>Verstraete B, De Block P, Robbrecht E (2025) A survey of generic names in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Gentianales</tp:taxon-name-part></tp:taxon-name>) with notes on context and patterns in naming. Taxon 74: 1153–1171. <ext-link xlink:href="10.1002/tax.13360" ext-link-type="doi">https://doi.org/10.1002/tax.13360</ext-link></mixed-citation>
      </ref>
      <ref id="B50">
        <mixed-citation>Weigend M (2002) Observations on the biogeography of the Amotape-Huancabamba Zone in northern Peru. In: Young K, Ulloa Ulloa C, Luteyn JL, Knapp S (Eds) Plant Evolution and Endemism in Andean South America. Botanical Review 68(1): 38–54. <ext-link xlink:href="https://www.jstor.org/stable/4354410" ext-link-type="uri">https://www.jstor.org/stable/4354410</ext-link> [accessed 28.01.2026]</mixed-citation>
      </ref>
      <ref id="B51">
        <mixed-citation>Weigend M (2004) Observaciones adicionales sobre la biogeografía de la zona de Amotape-Huancabamba en el norte del Perú: definiendo el límite suroriental. Revista Peruana de Biología 11(2): 127–134. <ext-link xlink:href="10.15381/rpb.v11i2.2447" ext-link-type="doi">https://doi.org/10.15381/rpb.v11i2.2447</ext-link></mixed-citation>
      </ref>
      <ref id="B52">
        <mixed-citation>Weigend M, Cano A, Rodríguez EF (2005) Nuevos registros y nuevas especies de la flora de la zona de AmotapeHuancabamba: Endemismos y límites biogeográficos. Revista Peruana de Biología 12(2): 249–274. <ext-link xlink:href="10.15381/rpb.v12i2.2398" ext-link-type="doi">https://doi.org/10.15381/rpb.v12i2.2398</ext-link></mixed-citation>
      </ref>
      <ref id="B53">
        <mixed-citation>Wolff D, Liede-Schumann S (2007) Evolution of flower morphology, pollen dimorphism, and nectar composition in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Arcytophyllum</tp:taxon-name-part></tp:taxon-name></italic>, a distylous genus of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Rubiaceae</tp:taxon-name-part></tp:taxon-name>. Organisms Diversity and Evolution 7(2): 106–123. <ext-link xlink:href="10.1016/j.ode.2006.02.005" ext-link-type="doi">https://doi.org/10.1016/j.ode.2006.02.005</ext-link></mixed-citation>
      </ref>
    </ref-list>
    <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.175775.suppl1</object-id>
        <object-id content-type="arpha">B7346F4B-2892-5B30-9595-E7EB69DEC439</object-id>
        <label>Supplementary material 1</label>
        <statement content-type="notes">
          <p>Taxon names with authors and GenBank accession numbers.</p>
        </statement>
        <media xlink:href="plecevo-159-244-s001.csv" mimetype="text" mime-subtype="csv" position="float" orientation="portrait" id="oo_1600434.csv">
          <uri content-type="original_file">https://binary.pensoft.net/file/1600434</uri>
        </media>
      </supplementary-material>
    </sec>
  </back>
</article>
