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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">103</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:77d0745d-c3a1-5248-81de-8cdc02bed84a</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:F56F6CF9-7502-4001-A751-35D5F2EF6CA0</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Arthropod Systematics &amp; Phylogeny</journal-title>
        <abbrev-journal-title xml:lang="en">ASP</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1863-7221</issn>
      <issn pub-type="epub">1864-8312</issn>
      <publisher>
        <publisher-name>Senckenberg Gesellschaft für Naturforschung</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/asp.84.e186815</article-id>
      <article-id pub-id-type="publisher-id">186815</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Chilopoda</subject>
          <subject>Myriapoda</subject>
          <subject>Notostigmophora</subject>
          <subject>Scutigeridae</subject>
          <subject>Scutigeromorpha</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Molecular systematics</subject>
          <subject>Morphology &amp; Anatomy</subject>
          <subject>Phylogeny</subject>
          <subject>Taxonomy</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Integrative morphological and molecular evidence reveals a new genus of scutigerid centipede from Hainan, China, with implications for its evolution and biogeography (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Scutigeromorpha">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Li</surname>
            <given-names>Qing</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0009-0008-6166-6021</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
          <role content-type="http://credit.niso.org/contributor-roles/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/methodology/">Methodology</role>
          <role content-type="http://credit.niso.org/contributor-roles/software/">Software</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Edgecombe</surname>
            <given-names>Gregory D.</given-names>
          </name>
          <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/supervision/">Supervision</role>
        </contrib>
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Jiang</surname>
            <given-names>Chao</given-names>
          </name>
          <email xlink:type="simple">jiangchao0411@126.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0003-1841-1169</uri>
          <xref ref-type="aff" rid="A1">1</xref>
          <role content-type="http://credit.niso.org/contributor-roles/conceptualization/">Conceptualization</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-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/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/supervision/">Supervision</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">State Key Laboratory for Quality Ensurance and Sustainable Use of Dao-di Herbs, National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences, Beijing, 100700, China</addr-line>
        <institution>Natural History Museum</institution>
        <addr-line content-type="city">London</addr-line>
        <country>United Kingdom</country>
        <uri content-type="ror">https://ror.org/039zvsn29</uri>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">The Natural History Museum, London SW7 5BD, United Kingdom</addr-line>
        <institution>China Academy of Chinese Medical Sciences</institution>
        <addr-line content-type="city">Beijing</addr-line>
        <country>China</country>
        <uri content-type="ror">https://ror.org/042pgcv68</uri>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Chao Jiang (<email xlink:type="simple">jiangchao0411@126.com</email>)</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2026</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>23</day>
        <month>06</month>
        <year>2026</year>
      </pub-date>
      <volume>84</volume>
      <fpage>447</fpage>
      <lpage>464</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/BE775A96-EFA0-59C5-A7C4-95482682CF8E">BE775A96-EFA0-59C5-A7C4-95482682CF8E</uri>
      <uri content-type="zoobank" xlink:href="https://zoobank.org/3E3F8CCB-E934-43D2-8748-01A75C247EFE">3E3F8CCB-E934-43D2-8748-01A75C247EFE</uri>
      <history>
        <date date-type="received">
          <day>30</day>
          <month>01</month>
          <year>2026</year>
        </date>
        <date date-type="accepted">
          <day>26</day>
          <month>04</month>
          <year>2026</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Qing Li, Gregory D. Edgecombe, Chao Jiang</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>
      <self-uri content-type="zoobank" xlink:type="simple">https://zoobank.org/3E3F8CCB-E934-43D2-8748-01A75C247EFE</self-uri>
      <abstract>
        <p>
          <bold>Abstract</bold>
        </p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov. et sp. nov</bold>. is described and illustrated based on specimens from Hainan Province, China. Morphological examination showed that these specimens belong to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> and share certain similarities with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905. Phylogenetic reconstruction based on five genes (nuclear 18S and 28S rRNA, mitochondrial 12S and 16S rRNA, and cytochrome <italic>c</italic> oxidase subunit I) indicated that the specimens form a distinct and well-supported clade that is sister group to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904, so a new genus is accordingly established. Combining the evolutionary history and biogeographic framework of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>, this study revealed the evolutionary significance of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. in the context of the Peninsular Indian Plate as a biotic ferry implicated in the origin of East and Southeast Asian lineages. Morphological similarities between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> are symplesiomorphies of a clade that includes these two genera, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904.</p>
      </abstract>
      <kwd-group>
        <label>Key words</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="class" reg="Chilopoda">Chilopoda</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>morphology</kwd>
        <kwd>new species</kwd>
        <kwd>phylogeny</kwd>
        <kwd>taxonomic key</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
      </kwd-group>
      <funding-group>
        <funding-statement>The research was supported by the CACMS Innovation Fund (nos. CI2024E003, nos. CI2024G00-09) and the Key Project at Central Government Level: the Ability Establishment of Sustainable Use for Valuable Chinese Medicine Resources (nos. 2060302).</funding-statement>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="1. Introduction" id="sec1">
      <title>1. Introduction</title>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Scutigeromorpha">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> is an ancient and morphologically highly specialized monophyletic group within the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class" reg="Chilopoda">Chilopoda</tp:taxon-name-part></tp:taxon-name>. It is taxonomically recognized as comprising three families (<xref ref-type="bibr" rid="B14">Edgecombe and Giribet 2009</xref>; <xref ref-type="bibr" rid="B15">Giribet and Edgecombe 2006</xref>, 2013). Among these, the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> exhibits the highest diversity, and is widely distributed across warm temperate, tropical and subtropical regions of the Americas, southern Europe and North/East Africa, and from Asia to Australia (<xref ref-type="bibr" rid="B11">Edgecombe 2011</xref>). Displaying a typical transoceanic disjunct distribution pattern, it serves as an ideal model for studying historical biogeography. <xref ref-type="bibr" rid="B28">Manivannan et al. (2024)</xref> confirmed that the scutigerid subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> originated in Gondwana by integrating molecular phylogenetics, fossil-calibrated time trees and biogeographic models. Subsequent dispersal events have shaped the group’s current distribution pattern centered on the Indo-Australian region, greatly advancing understanding of its macroevolutionary history.</p>
      <p>With the application of methodologies such as scanning electron microscopy, molecular phylogenetics, and biogeographic modeling, the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> is currently recognized to include approximately 17 genera (<xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>; <xref ref-type="bibr" rid="B11">Edgecombe 2011</xref>; <xref ref-type="bibr" rid="B3">Bonato et al. 2016</xref>; <xref ref-type="bibr" rid="B31">Porta and Giribet 2024</xref>). This comprises 14 valid genera and three monotypic genera of uncertain taxonomic status: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Diplacrophor">Diplacrophor</tp:taxon-name-part></tp:taxon-name></italic> Chamberlin, 1920 (Solomon Islands), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phanothereua">Phanothereua</tp:taxon-name-part></tp:taxon-name></italic> Chamberlin, 1958 (Solomon Islands), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuella">Thereuella</tp:taxon-name-part></tp:taxon-name></italic> Chamberlin, 1955 (Peru).</p>
      <p>In this study, an integrative taxonomic approach was employed to analyze a series of specimens collected from Hainan, China. Morphological examination indicates these specimens belong to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> and share several similarities with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905. Phylogenetic reconstruction based on five gene fragments (18S rRNA, 28S rRNA, 12S rRNA, 16S rRNA, and COI) reveals that these specimens form a distinct, well-supported clade, which is phylogenetically distinct from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> and the allied genera <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904. This pattern is consistent with the “Out-of-India” biogeographic model (<xref ref-type="bibr" rid="B28">Manivannan et al. 2024</xref>). Combining morphological and molecular evidence, we herein establish a new genus to accommodate these specimens.</p>
      <p>This paper provides a comprehensive morphological description and illustrations of the new genus and its type species. We present a diagnostic key to all known valid genera characterized by the absence of a pair of spine-bristles at the distal end of tarsus I (the traditionally delimited <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>). Based on molecular clock estimates and existing biogeographic frameworks (<xref ref-type="bibr" rid="B28">Manivannan et al. 2024</xref>), we further discuss the divergence time of this new lineage and its significance in regional evolution. New evidence is provided for improving the phylogenetic framework of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> and testing the colonization history of Gondwanan groups in Asia.</p>
    </sec>
    <sec sec-type="2. Material and methods" id="sec2">
      <title>2. Material and methods</title>
      <sec sec-type="2.1. Specimens, morphology, terminologies" id="sec3">
        <title>2.1. Specimens, morphology, terminologies</title>
        <p>Specimens were hand-collected and fixed in 75% ethanol. All specimens including the types of the newly described species are deposited in the National Resource Center for Chinese Materia Medica, China Academy of Chinese Medical Sciences (<named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>). Specimen identification numbers are in the format ‘<named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-YYYYMMDDXXX’, where <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content> is an abbreviation of the depository; YYYYMMDD is the date on which the specimen was collected; and XXX is a number given to this specimen according to the sequence of specimens collected on that date. Morphological terminology follows <xref ref-type="bibr" rid="B13">Edgecombe and Giribet (2006)</xref>, <xref ref-type="bibr" rid="B2">Bonato et al. (2010)</xref> and <xref ref-type="bibr" rid="B31">Porta and Giribet (2024)</xref>. Proportions of the female gonopods follow measurements described by <xref ref-type="bibr" rid="B39">Würmli (1973</xref>: fig. 1). Terminology for peristomatic structures (epipharynx and hypopharynx) and antennal trichomes and sensilla is as used by <xref ref-type="bibr" rid="B20">Koch and Edgecombe (2006)</xref> and <xref ref-type="bibr" rid="B33">Sombke et al. (2011)</xref>, respectively. Abbreviations and nomenclature used in this study are listed: T (singular) and TT (plural), stomatotergite(s); <abbrev xlink:title="Bayesian posterior probabilities">PP</abbrev>, Bayesian posterior probabilities; <abbrev xlink:title="Maximum likelihood bootstrap">BP</abbrev>, Maximum likelihood bootstrap.</p>
        <p>Specimens’ preservation and observation follow <xref ref-type="bibr" rid="B27">Li et al. (2026)</xref>. They were cleared in 70% lactic acid for observation of the venom glands. Morphological examination and imaging used a Leica DMC 6200 camera attached to a Leica M205 FA microscope and Keyence VHX7000N ultra-depth-of-field microscope. The habitus was photographed with a NIKKOR 105 mm macro lens attached to a Nikon D850. Electron micrographs were taken with a Hitachi S-3400N scanning electron microscope.</p>
        <p>To observe the hypopharynx and epipharynx, mouthparts were removed with tweezers and minute needles. The detached mouthparts (mandibles, first and second maxillae, and forcipules) were cleaned in an ultrasonic bath and fixed with 2.5% glutaraldehyde. The samples were then dehydrated through a graded ethanol series (50%, 70%, 80%, 90%, 95% for 20 min each, followed by three changes of 100% ethanol), then dehydrated ethanol was replaced with anhydrous tert-butanol, and the samples were freeze-dried in tert-butanol. The specimens were mounted on stainless steel stubs using double-sided conductive tape, and examined and electron micrographs taken under a Hitachi S-3400N scanning electron microscope.</p>
        <p>SKETCHBOOK 6.0.6 was used on an iPad 9 tablet to prepare line drawings from photographs. Grammarly was used to polish English in the manuscript, subsequently edited by the native-speaking coauthor.</p>
      </sec>
      <sec sec-type="2.2. Phylogenetic methods" id="sec4">
        <title>2.2. Phylogenetic methods</title>
        <p>Five markers (two nuclear ribosomal RNA genes: 18S and 28S rRNA; two mitochondrial ribosomal RNA genes: 12S and 16S rRNA; and the mitochondrial protein-encoding gene cytochrome <italic>c</italic> oxidase subunit I: COI) used in previous studies of scutigeromorph phylogeny (<xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>, <xref ref-type="bibr" rid="B14">2009</xref>; <xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>; <xref ref-type="bibr" rid="B16">Giribet and Edgecombe 2013</xref>; <xref ref-type="bibr" rid="B28">Manivannan et al. 2024</xref>; <xref ref-type="bibr" rid="B31">Porta and Giribet 2024</xref>; <xref ref-type="bibr" rid="B27">Ji et al. 2025</xref>; <xref ref-type="bibr" rid="B27">Li et al. 2026</xref>) were used to estimate the phylogenetic position of the new genus. Primers and PCR amplification procedures for specimens of the new genus and two additional species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> from China collected by us (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="chinensis">chinensis</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="kaijiangensis">kaijiangensis</tp:taxon-name-part></tp:taxon-name></italic><xref ref-type="bibr" rid="B27">Ji et al., 2025</xref>) follow <xref ref-type="bibr" rid="B16">Giribet and Edgecombe (2013)</xref>. Sample information, localities, and GenBank accession numbers are shown in Table S1.</p>
        <p>Representatives of three centipede orders outside <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Scutigeromorpha">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> were selected as outgroup taxa to root the tree. The data were analyzed using both Maximum Likelihood (<abbrev xlink:title="Maximum Likelihood">ML</abbrev>) and Bayesian Inference (<abbrev xlink:title="Bayesian Inference">BI</abbrev>). <abbrev xlink:title="Maximum Likelihood">ML</abbrev> analysis was performed on the IQ-TREE web tool on the Galaxy platform (<xref ref-type="bibr" rid="B30">Nguyen et al. 2015</xref>; <xref ref-type="bibr" rid="B34">Trifinopoulos et al. 2016</xref>; <xref ref-type="bibr" rid="B29">Minh 2020</xref>) and <abbrev xlink:title="Bayesian Inference">BI</abbrev> performed on the PhyloSuite V1.2.2 platform (<xref ref-type="bibr" rid="B43">Zhang et al. 2020</xref>). Sequences for each dataset were aligned using the FFT-NS-2 algorithm in MAFFT (<xref ref-type="bibr" rid="B21">Katoh and Standley 2013</xref>, <xref ref-type="bibr" rid="B22">2014</xref>; <xref ref-type="bibr" rid="B24">Katoh et al. 2005</xref>, <xref ref-type="bibr" rid="B23">2019</xref>). The resulting alignments were processed with Gblocks (<xref ref-type="bibr" rid="B6">Castresana 2000</xref>) under settings allowing gaps within the final blocks, applying less stringent constraints on flanking positions. The trimmed gene alignments were concatenated using the Concatenate datasets tool, with external Ns converted to “?”. The final concatenated alignment comprised five genes, further refined using Gblocks.</p>
        <p>The concatenated alignment was analyzed with the IQ-TREE web tool on the Galaxy platform. The best-fit nucleotide substitution model, selected by ModelFinder (<xref ref-type="bibr" rid="B19">Kalyaanamoorthy et al. 2017</xref>) under the Bayesian Information Criterion (<abbrev xlink:title="Bayesian Information Criterion">BIC</abbrev>), was SYM+I+R4. Branch support under the Maximum Likelihood (<abbrev xlink:title="Maximum Likelihood">ML</abbrev>) framework was calculated using the Ultrafast Jackknife method with 200,000 replicates (<xref ref-type="bibr" rid="B18">Hoang et al. 2018</xref>) and the Shimodaira–Hasegawa approximate likelihood ratio test (<abbrev xlink:title="Shimodaira–Hasegawa approximate likelihood ratio test">SH-aLRT</abbrev>) with 200,000 replicates (<xref ref-type="bibr" rid="B17">Guindon et al. 2010</xref>). Bayesian inference (<abbrev xlink:title="Bayesian Inference">BI</abbrev>) employed MRBAYES 3.2.6 (<xref ref-type="bibr" rid="B32">Ronquist et al. 2012</xref>) in the PhyloSuite v1.2.2 platform (<xref ref-type="bibr" rid="B43">Zhang et al. 2020</xref>), with 10,000,000 bootstrap replicates. SYM+I+G4 was chosen as the preferred model for <abbrev xlink:title="Bayesian Inference">BI</abbrev>, sampling every 1,000 generations and using 25% of the trees as burn-in. A split frequency of less than 0.01 was used to determine stationarity, and the consensus tree was constructed from the remaining trees. The resulting gene phylogenies were visualized in iTOL v5 (<xref ref-type="bibr" rid="B26">Letunic and Bork 2021</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="3. Results" id="sec5">
      <title>3. Results</title>
      <sec sec-type="3.1. Phylogenetic analyses" id="sec6">
        <title>3.1. Phylogenetic analyses</title>
        <p>Maximum Likelihood (<abbrev xlink:title="Maximum Likelihood">ML</abbrev>) and Bayesian Inference (<abbrev xlink:title="Bayesian Inference">BI</abbrev>) analyses based on concatenated sequences for the five genes recovered trees within which <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Scutigeromorpha">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> contained 80 and 60 strongly supported nodes (Maximum Likelihood bootstrap proportion (<abbrev xlink:title="Maximum likelihood bootstrap">BP</abbrev>) &gt; 90 or Bayesian posterior probabilities (<abbrev xlink:title="Bayesian posterior probabilities">PP</abbrev>) &gt; 95), respectively, with minor topological differences between them (Fig. <xref ref-type="fig" rid="F1">1</xref>).</p>
        <fig id="F1">
          <object-id content-type="doi">10.3897/asp.84.e186815.figure1</object-id>
          <object-id content-type="arpha">C6BED774-ACF4-55FD-A6E7-66D86D7BAEBD</object-id>
          <label>Figure 1.</label>
          <caption>
            <p>Maximum likelihood tree (left) and Bayesian consensus tree (right) based on 12S, 16S, 18S, 28S and COI sequences. <abbrev xlink:title="Maximum likelihood bootstrap">BP</abbrev> and <abbrev xlink:title="Bayesian posterior probabilities">PP</abbrev> values are shown next to each node, the support values with <abbrev xlink:title="Maximum likelihood bootstrap">BP</abbrev> &lt; 90 and <abbrev xlink:title="Bayesian posterior probabilities">PP</abbrev> &lt; 95 were not given.</p>
          </caption>
          <graphic xlink:href="arthropod-systematics-84-447-g001.jpg" id="oo_1690392.jpg">
            <uri content-type="original_file">https://binary.pensoft.net/fig/1690392</uri>
          </graphic>
        </fig>
        <p>Relationships between the three monophyletic scutigeromorph families are as in previous molecular phylogenies (<xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>, <xref ref-type="bibr" rid="B14">2009</xref>; <xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>; <xref ref-type="bibr" rid="B16">Giribet and Edgecombe 2013</xref>; <xref ref-type="bibr" rid="B1">Benavides et al. 2023</xref>; <xref ref-type="bibr" rid="B27">Li et al. 2026</xref>), with <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigerinidae">Scutigerinidae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> being sister groups to the exclusion of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Pselliodidae">Pselliodidae</tp:taxon-name-part></tp:taxon-name>. The <abbrev xlink:title="Maximum Likelihood">ML</abbrev> and <abbrev xlink:title="Bayesian Inference">BI</abbrev> trees both divided the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> into two clades, consistent with the subfamilial circumscriptions newly defined by <xref ref-type="bibr" rid="B27">Li et al. (2026)</xref>. In both trees, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> was divided into: (1) a clade comprising the genera <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tachythereua">Tachythereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dendrothereua">Dendrothereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1944, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Scutigera">Scutigera</tp:taxon-name-part></tp:taxon-name></italic> Lamarck, 1801; (2) a clade containing <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Sinothereua">Sinothereua</tp:taxon-name-part></tp:taxon-name></italic> Li, Edgecombe &amp; Jiang, 2026, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Edgethereua">Edgethereua</tp:taxon-name-part></tp:taxon-name></italic> Porta and Giribet, 2024, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lassophora">Lassophora</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ballonema">Ballonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904 and the traditionally recognized subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>.</p>
        <p>The species described herein was consistently and robustly nested within the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> clade, with strong support for a sister-group relationship with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> in both the <abbrev xlink:title="Maximum Likelihood">ML</abbrev> and <abbrev xlink:title="Bayesian Inference">BI</abbrev> trees. Based on the combined results, we support recognition of the Hainan species as a distinct new genus, which we hereby describe as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. Expanding the scope of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> to encompass <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">H.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. would require a rediagnosis that would not include the distinctive elongate spiculae that have long been used to diagnose <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B41">Würmli 1975a</xref>; <xref ref-type="bibr" rid="B10">Dyachkov 2026</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> group with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> in both <abbrev xlink:title="Maximum Likelihood">ML</abbrev> and <abbrev xlink:title="Bayesian Inference">BI</abbrev> trees, but the two methods differ in whether <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> are each other’s sister group (<abbrev xlink:title="Maximum Likelihood">ML</abbrev>) or if <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> is more closely related to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> + <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> (<abbrev xlink:title="Bayesian Inference">BI</abbrev>).</p>
      </sec>
      <sec sec-type="3.2. Taxonomy" id="sec7">
        <title>3.2. Taxonomy</title>
        <p>
          <bold>Order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order" reg="Scutigeromorpha">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> Pocock, 1895</bold>
        </p>
        <p>
          <bold>Family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> Leach, 1814</bold>
        </p>
        <p>
          <bold>Subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> Verhoeff, 1905</bold>
        </p>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom">Animalia</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>3.2.1.</label>
            <tp:taxon-name><object-id content-type="arpha">2C4D3141-C1F9-5EB9-9FE0-EB0A8C2E4DBD</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part>
                    	
                    		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/4E84E1DE-65A3-49FB-82EC-1C2530F0E96F</object-id>
                    	</tp:taxon-name>
            <tp:taxon-status>gen. nov.</tp:taxon-status>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Type species">
            <title>Type species.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. by monotypy and present designation.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Diagnosis">
            <title>Diagnosis.</title>
            <p>Antenna ca. 1.5 times as long as the body. First flagellum of antenna with 55–101 annulations (including node). Anterior projection of cephalic transverse sutures short, triangular, posterior part subparallel. Stomatotergites with a prominent white median longitudinal stripe; bristles (Stachelborsten) associated with elongate, conical, unpaired spines as long as bristles on TT4–7, and short paired spines on all tergites; the ratio of paired spines to unpaired spines associated with Stachelborsten on T6 is approximately 1:1 to 1:2, bristles (Stachelborsten) on borders with short paired spines along posterior margin. Elongate triangular spiculae, relatively sparse. Stomata elongate. Prefemur of legs 3–9 lacking a saw-like row of spines (sometimes legs 1–9). Legs 1–14 with pairs of tarsal papillae on consecutive tarsomeres, short-long alternation of tarsal papillae on legs 1–9, and uniform size of tarsal papillae on legs 10–14 (paired long papillae). Proarthron and metarthron of female gonopods subparallel-sided.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Etymology">
            <title>Etymology.</title>
            <p>The name <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> (feminine) is a compound derived from ‘<italic>Hainan</italic>-’ (Hainan Province, China) and the common scutigerid suffix -<italic>thereua</italic>, meaning “the scutigerid genus of Hainan Province, China”.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Remarks">
            <title>Remarks.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> in the cephalic sutures and the spines on the stomatotergites (all associated with a Stachelborste). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> differs from this genus in color, form of spiculae, and stronger Stachelborsten. The dark pigmented band on the stomatotergites of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> is rather diffuse, while there is a distinct white longitudinal stripe in the middle of the stomatotergites of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>. The spiculae of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> are short triangular or more elongate, relatively sparse. In marked contrast, the spiculae of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> are dense, long, needle-like and variably parallel-sided (<xref ref-type="bibr" rid="B35">Verhoeff 1904</xref>, <xref ref-type="bibr" rid="B36">1905</xref>; <xref ref-type="bibr" rid="B41">Würmli 1975a</xref>; <xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>: fig. 2f; <xref ref-type="bibr" rid="B11">Edgecombe 2011</xref>; <xref ref-type="bibr" rid="B10">Dyachkov 2026</xref>).</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilbarascutigera">Pilbarascutigera</tp:taxon-name-part></tp:taxon-name></italic> Edgecombe and Barrow, 2007 in the shape of its spiculae (<xref ref-type="bibr" rid="B12">Edgecombe 2007</xref>), but those of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> are relatively longer than in the other three genera. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> possesses a unique kinked cephalic suture, while that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilbarascutigera">Pilbarascutigera</tp:taxon-name-part></tp:taxon-name></italic> is of the M-shaped type common in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B11">Edgecombe 2011</xref>). Compared to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilbarascutigera">Pilbarascutigera</tp:taxon-name-part></tp:taxon-name></italic> with stoma-saddles weakly vaulted, the stoma-saddle of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> is moderately vaulted, while that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> is strongly vaulted and is approached by that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B12">Edgecombe and Barrow 2007</xref>). In addition, the stomatotergites bear spines associated with Tastborsten in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilbarascutigera">Pilbarascutigera</tp:taxon-name-part></tp:taxon-name></italic> (anterior tergites bearing Stachelborsten and Tastborsten but lacking spines), while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> bear Stachelborsten. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> in having the margins of the proarthron and mesarthron of the female gonopods diverging in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> v. subparallel-sided in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>; and the prefemur of legs 2–4 of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> lacking a saw-like row of spines whereas <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> lacks them on legs 3–9 (sometimes 1–9).</p>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. resembles <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allothereua">Allothereua</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parascutigera">Parascutigera</tp:taxon-name-part></tp:taxon-name></italic> in the spines on the stomatotergites being associated with Stachelborsten and the margin of the proarthron and mesarthron of female gonopods being subparallel-sided. The spiculae of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allothereua">Allothereua</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parascutigera">Parascutigera</tp:taxon-name-part></tp:taxon-name></italic> are, however, variably setiform (<xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>), rather than elongate triangular as in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>, and they are considerably denser in the former two genera. The Stachelborsten on TT5–7 of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parascutigera">Parascutigera</tp:taxon-name-part></tp:taxon-name></italic> are associated with paired spines, unpaired spines being wholly lacking (<xref ref-type="bibr" rid="B35">Verhoeff 1904</xref>, <xref ref-type="bibr" rid="B36">1905</xref>; <xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>; <xref ref-type="bibr" rid="B11">Edgecombe 2011</xref>), but both types of bristles/spines are present on the stomatotergites of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allothereua">Allothereua</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>.</p>
            <p>Like <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Seychellonema">Seychellonema</tp:taxon-name-part></tp:taxon-name></italic><xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. expresses a short-long alternation of tarsal papillae on successive tarsomeres. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Seychellonema">Seychellonema</tp:taxon-name-part></tp:taxon-name></italic>, this feature is present on legs 1–7, while legs 8–14 bear paired short papillae of uniform size (<xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>). However, in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>, the alternation is observed on legs 1–9, and uniform size of tarsal papillae on legs 10–14 (paired long papillae). <xref ref-type="bibr" rid="B42">Würmli (1975b)</xref> interpreted such alternation in the holotype of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lassophora">Lassophora</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="madagascariensis">madagascariensis</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905 as unreliable for generic diagnosis because the specimen is a praematurus, and mature specimens have uniform papillae. This morphology is, however, retained in mature specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Seychellonema">Seychellonema</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic>. Accordingly, we include it in the generic diagnosis.</p>
            <sec sec-type="3.2.2. Identification key" id="sec8">
              <title>3.2.2. Identification key</title>
              <p>What follows is a key to all known valid scutigerid genera characterized by the absence of a pair of spine-bristles at distal end of all tarsus I, with emphasis on tergal prominences.</p>
              <table-wrap content-type="key" position="anchor" orientation="portrait">
                <table>
                  <tbody>
                    <tr>
                      <td>
                        <bold>1</bold>
                      </td>
                      <td>Dorsal spine-bristle on prefemur of second maxillae; ventral spine-bristle lacking</td>
                      <td>
                        <bold>2</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>1’</bold>
                      </td>
                      <td>Dorsal and ventral spine-bristles on prefemur of second maxillae</td>
                      <td>
                        <bold>3</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>2</bold>
                      </td>
                      <td>Margins of proarthron and mesarthron of female gonopods diverging posteriorly</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Edgethereua">Edgethereua</tp:taxon-name-part></tp:taxon-name></italic> Porta and Giribet, 2024</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>2’</bold>
                      </td>
                      <td>Margins of proarthron and mesarthron of female gonopods weakly converging</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuoquima">Thereuoquima</tp:taxon-name-part></tp:taxon-name></italic> Bücherl, 1949</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>3</bold>
                      </td>
                      <td>Tergites bearing bristles and spines, lacking spiculae or spinulae</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tachythereua">Tachythereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>3’</bold>
                      </td>
                      <td>Tergites bearing bristles, spines and spiculae or spinulae</td>
                      <td>
                        <bold>4</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>4</bold>
                      </td>
                      <td>All tergites with spinulae</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Seychellonema">Seychellonema</tp:taxon-name-part></tp:taxon-name></italic> Butler, Edgecombe, Ball and Giribet, 2010</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>4’</bold>
                      </td>
                      <td>Tergites with simple spiculae</td>
                      <td>
                        <bold>5</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>5</bold>
                      </td>
                      <td>Spiculae as long as bristles, needle-like</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>5’</bold>
                      </td>
                      <td>Spiculae shorter than bristles</td>
                      <td>
                        <bold>6</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>6</bold>
                      </td>
                      <td>Tergites bearing few bristles/spines (spines lacking on stoma-saddles of TT6–7)</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Prionopodella">Prionopodella</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1925</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>6’</bold>
                      </td>
                      <td>Tergites bearing bristles associated with paired spines</td>
                      <td>
                        <bold>7</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>6’’</bold>
                      </td>
                      <td>Tergites bearing bristles associated with unpaired spines</td>
                      <td>
                        <bold>9</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>7</bold>
                      </td>
                      <td>Metarthron of female gonopods massive, hook-shaped</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pesvarus">Pesvarus</tp:taxon-name-part></tp:taxon-name></italic> Würmli, 1974</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>7’</bold>
                      </td>
                      <td>Metarthron of female gonopods slender, tapering distally</td>
                      <td>
                        <bold>8</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>8</bold>
                      </td>
                      <td>Short paired spines (no unpaired spines); margin of proarthron and mesarthron of female gonopods diverging posteriorly</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Prothereua">Prothereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1925</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>8’</bold>
                      </td>
                      <td>Variably long paired spines; margin of proarthron and mesarthron of female gonopods subparallel-sided</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Parascutigera">Parascutigera</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>9</bold>
                      </td>
                      <td>Anterior projection of cephalic sutures with divergent posterior part such that suture is kinked</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>9’</bold>
                      </td>
                      <td>Anterior projection of cephalic sutures subparallel</td>
                      <td>
                        <bold>10</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>10</bold>
                      </td>
                      <td>Tergites bearing isolated bristles (Tastborsten), few unpaired spines (spines lacking on stoma-saddles of TT6–7)</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Podothereua">Podothereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>10’</bold>
                      </td>
                      <td>Tergites bearing bristles (Tastborsten), unpaired spines (spines present on stoma-saddles of TT6–7)</td>
                      <td>
                        <bold>11</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>11</bold>
                      </td>
                      <td>Tergites bearing bristles (Tastborsten and Stachelborsten)</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilbarascutigera">Pilbarascutigera</tp:taxon-name-part></tp:taxon-name></italic> Edgecombe and Barrow, 2007</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>11’</bold>
                      </td>
                      <td>Tergites bearing bristles (Stachelborsten)</td>
                      <td>
                        <bold>12</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>12</bold>
                      </td>
                      <td>Margins of proarthron and mesarthron of female gonopods diverging posteriorly</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>12’</bold>
                      </td>
                      <td>Margins of proarthron and mesarthron of female gonopods subparallel-sided</td>
                      <td>
                        <bold>13</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>13</bold>
                      </td>
                      <td>Spiculae dense (at margins of most polygonal cuticular scales), variably setiform; uniform size of tarsal papillae on legs 1–14</td>
                      <td>
                        <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Allothereua">Allothereua</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905</bold>
                      </td>
                    </tr>
                    <tr>
                      <td>
                        <bold>13’</bold>
                      </td>
                      <td>Spiculae relatively sparse, elongate triangular; short-long alternation of tarsal papillae on legs 1–9</td>
                      <td><bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic> gen. nov</bold>.</td>
                    </tr>
                  </tbody>
                </table>
              </table-wrap>
            </sec>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom">Animalia</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>3.2.3.</label>
            <tp:taxon-name><object-id content-type="arpha">4A06CFEA-93C0-5B27-B13B-EC26A824AA74</object-id>
                    		<tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part>
                    	
                    		<object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/F4A46240-140E-43D6-AE9C-70594B6AFF8F</object-id>
                    	</tp:taxon-name>
            <tp:taxon-authority>gen. nov. et</tp:taxon-authority>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Material examined">
            <title>Material examined.</title>
            <p><bold>Holotype</bold>: CHINA • ♀; Mingfenggu Valley, Jianfengling, Ledong Li Autonomous County, Hainan Province, China (Fig. <xref ref-type="fig" rid="F2">2</xref>), <named-content content-type="dwc:verbatimCoordinates">18.7426°N 108.8400°E</named-content>, 990 m a.s.l., 22 June 2021, leg. Chao Jiang &amp; Tianyun Chen (labelled as <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210622177). <bold>Paratypes</bold>: CHINA • 2♀♀ 5♂♂, same as holotype, 22–23 June 2021, leg. Chao Jiang &amp; Tianyun Chen (<named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210622104, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210622174, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210622176, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210622179, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210623126, CCMI-20210623127, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210623128); CHINA • 2♀♀ 3♂♂, Mt. Diaoluoshan, Benhao Town, Lingshui Li Autonomous County, Hainan Province, China, <named-content content-type="dwc:verbatimCoordinates">18.7272°N 109.8724°E</named-content>, 900 m a.s.l., 24–25 June 2021, leg. Chao Jiang &amp; Tianyun Chen (<named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210624170, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210624171, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210625112, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210625173, <named-content content-type="dwc:institutional_code" xlink:title="Chinese Academy of Traditional Medicine" xlink:href="https://scientific-collections.gbif.org/institution/13b12b02-de2a-4fe2-b14f-303445b5d88f">CMMI</named-content>-20210625175).</p>
            <fig id="F2">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure2</object-id>
              <object-id content-type="arpha">6B112C91-BC05-5F68-9720-B576B4F18464</object-id>
              <label>Figure 2.</label>
              <caption>
                <p>Habitat of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>. (from type locality: Mingfenggu Valley, Jianfengling, Ledong Li Autonomous County, Hainan Province, China). <bold>A</bold> Panoramic view of type locality; <bold>B</bold> woodland on the mountain of type locality shown in <bold>A</bold>; <bold>C</bold> interior habitat of the woodland shown in <bold>B</bold>, the species was collected in relatively dry leaf litter or under stones; <bold>D</bold> individual of the species found under a stone; <bold>E</bold> live habitus of the species. <bold>A</bold>–<bold>D</bold> provided by Mr. Quanyu Ji, <bold>E</bold> provided by Mr. Jiazhou Lu.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g002.jpg" id="oo_1690393.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690393</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Diagnosis">
            <title>Diagnosis.</title>
            <p>As for genus.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Description">
            <title>Description.</title>
            <p>Body <bold>length</bold> (from anterior edge of head capsule to tip of gonopods): 18–23 mm in adult. — <bold>Colour</bold>: Head capsule brown (Fig. <xref ref-type="fig" rid="F3">3A, B</xref>); antennal base ferruginous grading distally to pale yellow; stomatotergites with a prominent white median longitudinal stripe, black and brown on both sides, margins dark brown (Figs <xref ref-type="fig" rid="F2">2D</xref>, <xref ref-type="fig" rid="F2">2E</xref>, <xref ref-type="fig" rid="F3">3C</xref>, <xref ref-type="fig" rid="F3">3D</xref>, <xref ref-type="fig" rid="F8">8A–F</xref>); stoma saddle red (Fig. <xref ref-type="fig" rid="F2">2D, E</xref>); legs yellow with conspicuous black bands on prefemur, femur, and tibia (Fig. <xref ref-type="fig" rid="F2">2D, E</xref>, 3A); sternites pale yellow; female gonopods yellow (Fig. <xref ref-type="fig" rid="F8">8G</xref>) (based on living specimens). — <bold>Head capsule</bold>: Anterior projection of the cephalic transverse sutures short, triangular, posterior part parallel; transverse suture long; longitudinal median depression extends from the base of the antennae to between the two eyes (Figs <xref ref-type="fig" rid="F3">3A</xref>, <xref ref-type="fig" rid="F3">3B</xref>, <xref ref-type="fig" rid="F10">10A</xref>, <xref ref-type="fig" rid="F10">10C</xref>). Antenna ca. 1.5 times as long as body length. First flagellum of antenna with 55–101 annulations (including the first node). Annulations much wider than long (Figs <xref ref-type="fig" rid="F3">3A</xref>, <xref ref-type="fig" rid="F3">3B</xref>, <xref ref-type="fig" rid="F10">10A</xref>, <xref ref-type="fig" rid="F10">10C</xref>), with three or four whorls of flattened trichomes, a few beak-like sensilla, and a few setae (trichoid sensilla) associated with paired spines connected to the base form a single whorl encircling the distal end of the annulation. Tömösváry’s organ small (Fig. <xref ref-type="fig" rid="F3">3B</xref>), in typical position between eye and base of antenna (Fig. <xref ref-type="fig" rid="F2">2D, E</xref>). — <bold>Epipharynx</bold>: Arrangement typical for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family" reg="Scutigeridae">Scutigeridae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B20">Koch and Edgecombe 2006</xref>: fig. 1) (Fig. <xref ref-type="fig" rid="F4">4</xref>). Lateral bar of labral trapezoid with narrow longitudinal groove along whole length of bar (Fig. <xref ref-type="fig" rid="F4">4A</xref>). Labral bristles differentiated into narrow outer band of short, pectinate bristles and wider inner band of longer simple bristles (Fig. <xref ref-type="fig" rid="F4">4A, C</xref>). Two clusters of sensilla along mid-line of labral trapezoid: more distal unpaired, transverse group of a few dozen bottle-shaped sensilla at termination of median ridge (Fig. <xref ref-type="fig" rid="F4">4B</xref>); the three proximal sensilla clusters of the labral part are composed of nipple-and bottle-shaped sensilla, median sensilla completely distal to lateral clusters (Fig. <xref ref-type="fig" rid="F4">4D</xref>). Chevron-shaped spine row of triangular and distally-curved denticles at the border (Fig. <xref ref-type="fig" rid="F4">4A, E, F</xref>) between labral and clypeal part of epipharynx; immediately proximal to the spine row is a rhomboid branched-spine field having transverse axis approximately 1.5 times the length of longitudinal axis, with a dense median field of pectinate spines (Fig. <xref ref-type="fig" rid="F4">4E</xref>). Broadly ovate cluster of bottle-shaped sensilla on medial part of clypeal triangle, a short distance behind dense field of branching spines (Fig. <xref ref-type="fig" rid="F4">4G</xref>). Lateral clusters of nipple-shaped sensilla positioned in a depression within the non-sclerotized area distal to the oblique bar and proximal to the edge extending from the submarginal armature to the transverse bar of the labral trapezoid. — <bold>Hypopharynx</bold>: Elongate, deeply projecting into preoral chamber (Fig. <xref ref-type="fig" rid="F5">5</xref>). Bars of lateral sclerotised fork with lateral bulges. Median excavation on proximal part of frontal and distal part of hypopharynx surface bordered laterally by pectinate bristles (Fig. <xref ref-type="fig" rid="F5">5A, D</xref>). Proximal cluster of sensilla (Fig. <xref ref-type="fig" rid="F5">5B, C</xref>) composed of a few sparse elements distributed in rows between mouth and a median elevation proximal to the median excavation. Area between the median excavation and the converging flattened bars of the proximal fork is sub-triangular with serrated spines at the proximal end of the carina (Fig. <xref ref-type="fig" rid="F5">5B, D</xref>). Inner margin of the tongue tip has strong irregular folds (Fig. <xref ref-type="fig" rid="F5">5E–G</xref>), on which nipple-shaped sensilla are distributed (Fig. <xref ref-type="fig" rid="F5">5E–G</xref>). — <bold>Mandible</bold>: Incisor region composed of three teeth, all with surface smooth and three cusps. Approximately 16 pectinate lamellae in gnathal lobe. Molar plate surrounded by a Haarpolster of approximately the same length. (Fig. <xref ref-type="fig" rid="F6">6A</xref>) — <bold>Second maxilla</bold>: Dorsal and ventral spine bristles on prefemur, four spine bristles on femur; two dorsal spine bristles on tibia (Fig. <xref ref-type="fig" rid="F6">6B</xref>). Ventral side of all podomeres and dorsal side of femur with rows of bristles. All spine bristles with similar surface details, spiniform scales, short proximally, becoming more elongate distally to confer a ridged or fluted surface to distal third of spine bristle. — <bold>Forcipular segment</bold>: Four spine bristles on anterior margin of coxa approximately equal in length to coxa. Spine comb of tarsus covering the entire inner side of the segment and composed of flattened setae mostly inserted by pairs in the bases and terminating in a rounded bulb. Ventrolateral side of trochanteroprefemur and femur with rows of spine bristles. Venom gland extending to distal part of tibia (Fig. <xref ref-type="fig" rid="F6">6C</xref>). — <bold>Stomatotergites</bold>: Stomatotergites with rugose surface (Fig. <xref ref-type="fig" rid="F7">7</xref>), posterior borders evenly rounded (Figs <xref ref-type="fig" rid="F2">2E</xref>, <xref ref-type="fig" rid="F3">3A</xref>, <xref ref-type="fig" rid="F3">3C</xref>, <xref ref-type="fig" rid="F3">3D</xref>, <xref ref-type="fig" rid="F7">7C</xref>, <xref ref-type="fig" rid="F7">7G</xref>, <xref ref-type="fig" rid="F8">8A–F</xref>, <xref ref-type="fig" rid="F10">10A</xref>, <xref ref-type="fig" rid="F10">10B</xref>). Stoma saddles moderately inflated; stomata elongate; with stomata 0.5–0.7 times the length of the stoma saddles (Fig. <xref ref-type="fig" rid="F7">7C, D</xref>). Stomatotergites bearing bristles (Stachelborsten) associated with elongate, conical, unpaired spines, as long as the bristles, or short paired spines (Figs <xref ref-type="fig" rid="F7">7A–E</xref>, <xref ref-type="fig" rid="F10">10F</xref>); spiculae short triangular or more elongate, emerging between the scutes, each separated by several polygonal scales that lack spiculae, relatively sparse (Figs <xref ref-type="fig" rid="F7">7</xref>, <xref ref-type="fig" rid="F10">10F</xref>); spinulae occur exclusively on T8 (Fig. <xref ref-type="fig" rid="F7">7G, H</xref>). Unpaired spines usually initially occur on T4 (on TT4–7), sparse, and always associated with a Stachelborste; 27–51 bristles associated with a spine on T6 (Figs <xref ref-type="fig" rid="F8">8D</xref>, <xref ref-type="fig" rid="F10">10D</xref>), uniformly distributed. Paired spines short; association with Stachelborste on TT1–8, 30–55 bristles associated with spines on T6 (Figs <xref ref-type="fig" rid="F8">8D</xref>, <xref ref-type="fig" rid="F10">10D</xref>). In adults, the ratio of paired spines to unpaired spines associated with Stachelborsten on T6 is approximately 1:1 to 1:2. Stomatotergite lateral margins and posterior margin with marginal bristles (Stachelborsten) with short, conical unpaired spines (Figs <xref ref-type="fig" rid="F7">7F</xref>, <xref ref-type="fig" rid="F10">10G</xref>). — <bold>Legs</bold>: Tarsus I and II tarsomeres as follow (range in tarsus I/tarsus II): leg 1, 11–16/25–31; leg 2, 10–11/27–30; leg 3, 10–11/25–30; leg 4, 7–9/24–30; leg 5, 7–9/24–30; leg 6, 7–9/24–27; leg 7, 7–8/19–29; leg 8, 6–7/22–27; leg 9, 6–8/24–28; leg 10, 6–7/28–29; leg 11, 6–8/29–30; leg 12, 6–8/26–32: leg 13, 8/31–34; leg 14, 7–9/29–32. Prefemoral spine-bristles in a 2/1 pattern on legs 1–14; femoral spine-bristles 1/2 on legs 1–14; tibial spine-bristles 1/1 on legs 1–11 and 1/2 on legs 12–14. Prefemur of legs 3–9 lacking a saw-like row of spines (sometimes legs 1–9). Pair of spine-bristles lacking at distal end of tarsus I (Fig. <xref ref-type="fig" rid="F10">10I</xref>). Pairs of tarsal papillae on legs 1–14, expression of short-long alternation of tarsal papillae on successive tarsomeres on legs 1–9 (sometimes legs 1–10) (Figs <xref ref-type="fig" rid="F6">6D</xref>, <xref ref-type="fig" rid="F9">9A–D</xref>, <xref ref-type="fig" rid="F10">10J</xref>); legs 10–14 (or legs 11–14) with paired long tarsal papillae (Figs <xref ref-type="fig" rid="F6">6E</xref>, <xref ref-type="fig" rid="F9">9E–H</xref>, <xref ref-type="fig" rid="F10">10K</xref>). Setal cluster associated with tarsal papillae sparse, arranged in longitudinal bands of 2–3 setae. Resilient sole hairs originate near posteromedial edge of tarsal papillae, extending to approximately one-third length of the succeeding tarsomere. — <bold>Sternites</bold>: Sternites with median swelling. Longitudinal median furrow visible, extending two-thirds length of the sternite. All sternites are scattered with setae, no hairs, spines or spiculae. — <bold>Gonopods</bold>: Female gonopods with maximum length 1.5–2.2 times maximum width; longitudinal median suture in proarthron complete (Figs <xref ref-type="fig" rid="F8">8G</xref>, <xref ref-type="fig" rid="F9">9I</xref>, <xref ref-type="fig" rid="F10">10E</xref>). Lateral margins of proarthron nearly parallel (lateral margins of the proarthron of the juvenile gonopods diverge posteriorly). Subtriangular depression on proarthron deep, lacking setae. Proarthron 1–1.5 times length of mesarthron (Figs <xref ref-type="fig" rid="F8">8G</xref>, <xref ref-type="fig" rid="F9">9I</xref>, <xref ref-type="fig" rid="F10">10E</xref>). Coarsest setae on proarthron slightly thicker than those on mesarthron apart from cluster at distomedial corner of mesarthron, this cluster usually composed of 6–10 setae (Fig. <xref ref-type="fig" rid="F9">9J, K</xref>). Sinus between mesarthron usually broadly parabolic, its apex typically weakly pointed (Fig. <xref ref-type="fig" rid="F9">9J</xref>). Width of mesarthron 0.4–0.8 times maximum width of sinus. Proarthron + mesarthron 1.7–2.2 times length of metarthron. Outer margin of metarthron arched (Fig. <xref ref-type="fig" rid="F9">9L</xref>). Ventral surface of metarthron scattered with small, pointed conical sensilla; outer margin of the metarthron bears 15–16 sensilla coeloconica, arranged roughly in a single row (Figs <xref ref-type="fig" rid="F9">9L</xref>, <xref ref-type="fig" rid="F9">9M</xref>, <xref ref-type="fig" rid="F10">10H</xref>). A few setae on dorsal side of metarthron (Figs <xref ref-type="fig" rid="F9">9L</xref>, <xref ref-type="fig" rid="F10">10E</xref>). Subanal plate drop-shaped; smooth, seta-free band in the middle of the subanal plate; setae present on outer surface of subanal plate (Fig. <xref ref-type="fig" rid="F8">8G, H</xref>). Both pairs of styliform male gonopods densely covered with setae, terminal part with short, spiniform setae between longer setae (Fig. <xref ref-type="fig" rid="F8">8H</xref>).</p>
            <fig id="F3">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure3</object-id>
              <object-id content-type="arpha">57FD5003-8D41-5656-B559-13FA367BE3B6</object-id>
              <label>Figure 3.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., holotype. <bold>A</bold> Habitus, dorsal view, body length 23 mm; <bold>B</bold> head, dorsal view; <bold>C</bold> T1, dorsal view; <bold>D</bold> T2, dorsal view. Abbreviations: lmd, longitudinal median depression; ap, anterior projection of the (cephalic) transverse suture; To, Tömösváry’s organ; t, (cephalic) transverse suture; ce, compound eye.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g003.jpg" id="oo_1690394.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690394</uri>
              </graphic>
            </fig>
            <fig id="F4">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure4</object-id>
              <object-id content-type="arpha">BF22390C-CB4D-58F1-84EB-EDB05CE05DD8</object-id>
              <label>Figure 4.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., paratypes. <bold>A</bold> Internal view of epipharynx, the labels *<bold>B</bold>–*<bold>G</bold> indicate the corresponding positions shown in panels <bold>B</bold>–<bold>G</bold>, and the lateral orientation is marked; <bold>B</bold> detail of <bold>A</bold>, distal part of the epipharynx, distal cluster of sensilla of the medial labral part; <bold>C</bold> lateral (left) part of the epipharynx, labral bristles, present on both sides; <bold>D</bold> detail of <bold>A</bold>, proximal part of the epipharynx, proximal cluster of sensilla of the medial labral part; <bold>E</bold> detail of <bold>A</bold>, field of branching spines on clypeal part of epipharynx; <bold>F</bold> detail of <bold>E</bold>, showing the chevron-shaped spine row; <bold>G</bold> sensilla proximal to dense field of branching spines on clypeal triangle, corresponding to the proximal part of the structure shown in panel <bold>E</bold>; <bold>H</bold> detail of <bold>A</bold>, sensilla on lateral (right) part of epipharynx, present on both sides, situated within a depression.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g004.jpg" id="oo_1690395.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690395</uri>
              </graphic>
            </fig>
            <fig id="F5">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure5</object-id>
              <object-id content-type="arpha">2C6DDFF7-A4C0-5314-8071-BAA15332D009</object-id>
              <label>Figure 5.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., paratype. <bold>A</bold> Frontal view of hypopharynx, the labels *<bold>B</bold>, *<bold>D</bold>, and *<bold>E</bold> indicate the corresponding positions shown in panels <bold>B</bold>, <bold>D</bold>, and <bold>E</bold> respectively, and the proximal and distal orientations are also marked; <bold>B</bold> detail of <bold>A</bold>, proximal part of the hypopharynx, mouth and area between the median excavation and the converging flattened bars of the proximal fork; <bold>C</bold> detail of <bold>B</bold>, showing sensilla; <bold>D</bold> detail of <bold>A</bold>, flattened bristles of proximal part of excavation; <bold>E</bold> distal end of hypopharynx, showing flattened bristles and sensilla, the labels *<bold>F</bold> and *<bold>G</bold> indicate the corresponding positions shown in panels <bold>F</bold> and <bold>G</bold>; <bold>F</bold> detail of <bold>E</bold>, showing nipple-shaped sensilla; <bold>G</bold> detail of <bold>E</bold>, showing pectinate bristles on distal portion.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g005.jpg" id="oo_1690396.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690396</uri>
              </graphic>
            </fig>
            <fig id="F6">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure6</object-id>
              <object-id content-type="arpha">B15BB974-8DF4-5F54-A296-D119D63A275A</object-id>
              <label>Figure 6.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., paratype. <bold>A</bold> Mandible, internal view; <bold>B</bold> second maxilla (left), prolateral view; <bold>C</bold> part of the forcipular segment, ventral view; <bold>D</bold> tarsal papillae on tarsomeres of leg 8 tarsus II, posterior view, showing the short-long alternation of tarsal papillae; <bold>E</bold> tarsal papillae on tarsomeres of leg 10 tarsus II, posterior view, showing long papillae of consistent size. Triangles represent long papillae, and circles represent short papillae.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g006.jpg" id="oo_1690397.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690397</uri>
              </graphic>
            </fig>
            <fig id="F7">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure7</object-id>
              <object-id content-type="arpha">16A804DE-7549-52E8-82EC-FB05F1E2DAE7</object-id>
              <label>Figure 7.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., paratype. <bold>A</bold> T2, dorsal view; <bold>B</bold> T5, dorsal view. <bold>C</bold> T6, dorsal view; <bold>D</bold> stoma saddles of T6, dorsal view; <bold>E</bold> bristles (Stachelborsten), spines, spiculae on T6, dorsal view; <bold>F</bold> paired spines with bristle on margin of T6, dorsal view. <bold>G</bold> T8, dorsal view; <bold>H</bold> detail of <bold>G</bold>, showing spiculae and spinulae on T8, the spiculae clearly emerging between the scutes, dorsal view. Abbreviations: St, Stachelborsten; p.s., paired spines; s, spine; sc, spiculae; sn, spinulae.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g007.jpg" id="oo_1690398.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690398</uri>
              </graphic>
            </fig>
            <fig id="F8">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure8</object-id>
              <object-id content-type="arpha">A92E6183-55FC-5524-A244-3B32F535F1C9</object-id>
              <label>Figure 8.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., <bold>A</bold>–<bold>G</bold> holotype, <bold>H</bold> paratype. <bold>A</bold> T3, dorsal view; <bold>B</bold> T4, dorsal view; <bold>C</bold> T5, dorsal view; <bold>D</bold> T6, dorsal view; <bold>E</bold> T7, dorsal view; <bold>F</bold> T8, (right-)dorsal view; <bold>G</bold> postpedal segments, including female gonopods and subanal plate, with relevant structures labeled, left-ventral view; <bold>H</bold> postpedal segments, including male gonopods, left-ventral view.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g008.jpg" id="oo_1690399.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690399</uri>
              </graphic>
            </fig>
            <fig id="F9">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure9</object-id>
              <object-id content-type="arpha">150DE0B0-97CA-5E66-99F9-5E157219FBA3</object-id>
              <label>Figure 9.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., paratype. <bold>A</bold> Tarsus II of leg 6, posterior view; <bold>B</bold> detail of <bold>A</bold>, tarsal papillae on tarsomeres of leg 6 tarsus II, lateral view; <bold>C</bold> detail of <bold>B</bold>, showing long papillae of consistent size, lateral view; <bold>D</bold> detail of <bold>B</bold>, showing short papillae of consistent size, lateral view; <bold>E</bold> tarsus II of leg 12, posterior view; <bold>F</bold> detail of <bold>E</bold>, tarsal papillae on tarsomeres of leg 12 tarsus II, lateral view; <bold>G</bold>, <bold>H</bold> detail of <bold>F</bold>, showing long papillae of consistent size, lateral view; <bold>I</bold> female gonopods and subanal plate, ventral view; <bold>J</bold> detail of <bold>I</bold>, showing mesarthron, ventral view; <bold>K</bold> detail of <bold>I</bold>, distal end of mesarthron, ventral view; <bold>L</bold> detail of <bold>I</bold>, showing metarthron, ventral view; <bold>M</bold> detail of <bold>L</bold>, showing sensillum on metarthron, ventral view.</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g009.jpg" id="oo_1690400.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690400</uri>
              </graphic>
            </fig>
            <fig id="F10">
              <object-id content-type="doi">10.3897/asp.84.e186815.figure10</object-id>
              <object-id content-type="arpha">2F4E4FF1-2F46-59ED-A6FE-2F8BAAD7DA63</object-id>
              <label>Figure 10.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="albilineata">albilineata</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov</bold>., <bold>A</bold>–<bold>E</bold> and <bold>I</bold> holotype, <bold>F</bold>–<bold>H</bold> and <bold>J</bold>–<bold>K</bold> paratype. <bold>A</bold>, <bold>B</bold> Habitus, showing head and TT1–8, dorsal view; <bold>C</bold> head, dorsal view; <bold>D</bold> T6, dorsal view; <bold>E</bold> female gonopods, (right-)ventral view; <bold>F</bold> detail of T6, dorsal view; <bold>G</bold> detail of posterior border of T6, right side, dorsal view; <bold>H</bold> right metarthron of female gonopods, dorsal view; <bold>I</bold> leg 6 (left) from prefemur to claw, posterior view; <bold>J</bold> detail of tarsal papillae on tarsus II of leg 6, lateral view; <bold>K</bold> detail of tarsal papillae on tarsus II of leg 10, lateral view. Triangles (<bold>J</bold>, <bold>K</bold>) represent long papillae, circles (<bold>J</bold>) represent short papillae. Scale bars: 500 μm (<bold>A</bold>–<bold>E</bold>, <bold>I</bold>); 300 μm (<bold>F</bold>, <bold>H</bold>); 100 μm (<bold>G</bold>, <bold>J</bold>, <bold>K</bold>).</p>
              </caption>
              <graphic xlink:href="arthropod-systematics-84-447-g010.jpg" id="oo_1690401.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1690401</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Distribution">
            <title>Distribution.</title>
            <p>China (Hainan).</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Etymology">
            <title>Etymology.</title>
            <p>The specific epithet albilineata is a Latin feminine adjective. It is a compound formed from “albi-”, derived from the stem of the Latin adjective <italic>albus</italic>, meaning “white”, and “-lineata”, the feminine past participle of the Latin verb <italic>lineare</italic> (to draw a line), meaning “lined” or “striped.” The name means “white-striped,” describing the distinct white longitudinal stripe in the middle of the tergites of this species.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
    <sec sec-type="4. Discussion" id="sec9">
      <title>4. Discussion</title>
      <p>Scutigeromorphs in the Asia-Australia region are highly diverse, with morphological similarity between species, making identification difficult (<xref ref-type="bibr" rid="B12">Edgecombe and Barrow 2007</xref>). Phylogenetic analyses show that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. is closely related to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic>, but there are significant morphological differences between the two genera. Although <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. is morphologically similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic>, they can be distinguished by the shape of margins of the proarthron and mesarthron of female gonopods and the saw-like row of spines of the prefemur of different ranges of legs. In addition, molecular phylogenetic analysis strongly supports <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. as an independent evolutionary clade. Based on a combination of diagnostic morphological features and molecular evidence, we classify the described specimens as a new genus.</p>
      <p>Furthermore, based on the phylogenetic trees constructed in this study, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Edgethereua">Edgethereua</tp:taxon-name-part></tp:taxon-name></italic> is stably positioned within the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>. This differs from the findings of <xref ref-type="bibr" rid="B31">Porta and Giribet (2024)</xref> and <xref ref-type="bibr" rid="B27">Li et al. (2026)</xref>, which reported conflicting results between the <abbrev xlink:title="Maximum Likelihood">ML</abbrev> and <abbrev xlink:title="Bayesian Inference">BI</abbrev> analyses. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lassophora">Lassophora</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905, from Madagascar and Mozambique, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ballonema">Ballonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904, from New Guinea, are here allied to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>, instead of forming a clade or grade excluded from either of the subfamilies <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Scutigerinae">Scutigerinae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>; <xref ref-type="bibr" rid="B16">Giribet and Edgecombe 2013</xref>; <xref ref-type="bibr" rid="B31">Porta and Giribet 2024</xref>). This result confirms that the traditional <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Scutigerinae">Scutigerinae</tp:taxon-name-part></tp:taxon-name> were reciprocally paraphyletic groups (<xref ref-type="bibr" rid="B13">Edgecombe and Giribet 2006</xref>; <xref ref-type="bibr" rid="B14">Edgecombe and Giribet 2009</xref>; <xref ref-type="bibr" rid="B4">Butler et al. 2010</xref>; <xref ref-type="bibr" rid="B16">Giribet and Edgecombe 2013</xref>; <xref ref-type="bibr" rid="B27">Li et al. 2026</xref>), indicating that relying only on a single morphological character (presence or absence of tarsus I spine bristles) is insufficient to reflect the true evolutionary history and prompting us to re-examine the early evolution and biogeographic patterns of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>.</p>
      <p>The biogeographic reconstruction by <xref ref-type="bibr" rid="B28">Manivannan et al. (2024)</xref> indicates that the ancestors of scutigeromorphs were widely distributed on Gondwana, and their major early lineage diversification occurred before the complete isolation of the main continental blocks. Following <xref ref-type="bibr" rid="B27">Li et al. (2026)</xref>, the affinities of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lassophora">Lassophora</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ballonema">Ballonema</tp:taxon-name-part></tp:taxon-name></italic> to the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> allow the effective divergence time of this subfamily to be traced back to at least the Early Cretaceous, around 145 million years ago (= mya). The transoceanic dispersal events through which <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lassophora">Lassophora</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Ballonema">Ballonema</tp:taxon-name-part></tp:taxon-name></italic> originated at this time may be more reasonably explained as an early divergence within the widely distributed ancestral group of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name>, rather than strictly corresponding to a specific continental breakup event. This conclusion differs from the view in <xref ref-type="bibr" rid="B16">Giribet and Edgecombe (2013)</xref>, which preferred to explain the phylogenetic relationships of this clade by vicariance during the Early Jurassic to Early Cretaceous. The ancestral group of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> had already achieved initial diversification on the drifting Indian Plate during the early to middle Cretaceous.</p>
      <p>Estimates based on the molecular clock (<xref ref-type="bibr" rid="B28">Manivannan et al. 2024</xref>) suggest that the likely divergence time of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. is comparable to that of its sister group, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic>, but slightly later than that of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> (the Indian diversification of which dates to around 78 mya), likely occurring between 60 and 72 mya. This temporal sequence is highly consistent with the geological history of the Peninsular Indian Plate. The ancestral divergence of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> occurred in the late stage of the isolated drift of the Indian Plate. In contrast, the divergence between the new genus and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> is inferred to have occurred during the critical window of the initial collision between the Indian Plate and the Eurasian Plate (about 60 to 50 mya). It is inferred that the ancestors of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. were pioneer groups that dispersed northward around the time of the plate collision. They have evolved independently to this day due to geographical isolation. The morphological similarity between <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Hainanthereua">Hainanthereua</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. nov</bold>. and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopodina">Thereuopodina</tp:taxon-name-part></tp:taxon-name></italic> originates from their common ancestor on the Indian Plate during the Cretaceous Period, the similarities between these two genera corresponding to symplesiomorphies of the clade that unites them with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuonema">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Thereuopoda">Thereuopoda</tp:taxon-name-part></tp:taxon-name></italic>. This also indicates that the diversification within Asian lineages is more complex than previously thought, which may involve multiple dispersal and vicariance events toward the edge of East Asia.</p>
      <p>The discovery of the new genus provides key empirical evidence from East Asia for testing and refining the “Indian Plate as a biological ferry” hypothesis. It also extends the evolutionary history of endemic Asian <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily" reg="Thereuoneminae">Thereuoneminae</tp:taxon-name-part></tp:taxon-name> to older geological periods, laying a more solid foundation for understanding the global distribution pattern of this ancient group. Future research should conduct more systematic surveys of scutigeromorph species in Southeast Asia and southern China to trace the complete trajectory of this “northward dispersal” route.</p>
    </sec>
    <sec sec-type="5. Declarations" id="sec10">
      <title>5. Declarations</title>
      <p><bold>Authors’ contributions</bold>. Qing Li: Conceptualization, Methodology, Software, Data Curation, Writing − Original Draft, Writing − Review &amp; Editing. Gregory D. Edgecombe: Data Curation, Supervision, Writing − Review &amp; Editing. Chao Jiang: Conceptualization, Resources, Data Curation, Methodology, Supervision, Writing − Review &amp; Editing, Project administration, Funding acquisition.</p>
      <p><bold>Conflict of interest</bold>. The authors declare that there is no conflict of interest.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>6. Acknowledgements</title>
      <p>We sincerely thank editors Martin Schwentner, Klaus-Dieter Klass, reviewer Andy Sombke and anonymous reviewers for reviewing the manuscript and providing valuable comments. We are grateful to Mr. Jiazhou Lu (Shaanxi Vocational Academy of Art, China) and Mr. Quanyu Ji (Hebei University, China) for providing habitat photos of the new species. We also thank Mr. Feiyu Huang (Northeast Forestry University, China) for help with the molecular work. The research was supported by the CACMS Innovation Fund (nos. CI2024E003, nos. CI2024G00-09) and the Key Project at Central Government Level: the Ability Establishment of Sustainable Use for Valuable Chinese Medicine Resources (nos. 2060302).</p>
    </ack>
    <ref-list>
      <title>7. References</title>
      <ref id="B1">
        <mixed-citation>Benavides LR, Edgecombe GD, Giribet G (2023) Re-evaluating and dating myriapod diversification with phylogenomics under a regime of dense taxon sampling. Molecular Phylogenetics and Evolution 178: 107621. <ext-link xlink:href="10.1016/j.ympev.2022.107621" ext-link-type="doi">https://doi.org/10.1016/j.ympev.2022.107621</ext-link></mixed-citation>
      </ref>
      <ref id="B2">
        <mixed-citation>Bonato L, Edgecombe GD, Lewis JGE, Minelli A, Pereira LA, Shelley RM, Zapparoli M (2010) A common terminology for the external anatomy of centipedes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>). Zookeys 69: 17–51. <ext-link xlink:href="10.3897/zookeys.69.737" ext-link-type="doi">https://doi.org/10.3897/zookeys.69.737</ext-link></mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation>Bonato L, Chagas Júnior A, Edgecombe GD, Lewis JGE, Minelli A, Pereira LA, Shelley RM, Stoev P, Zapparoli M (2016) ChiloBase 2.0 – A World Catalogue of Centipedes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>). <ext-link xlink:href="https://chilobase.biologia.unipd.it" ext-link-type="uri">https://chilobase.biologia.unipd.it</ext-link></mixed-citation>
      </ref>
      <ref id="B4">
        <mixed-citation>Butler AD, Edgecombe GD, Ball AD, Giribet G (2010) Resolving the phylogenetic position of enigmatic New Guinea and Seychelles <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>): A molecular and morphological assessment of Ballonemini. Invertebrate Systematics 24: 539–559. <ext-link xlink:href="10.1071/IS10037" ext-link-type="doi">https://doi.org/10.1071/IS10037</ext-link></mixed-citation>
      </ref>
      <ref id="B5">
        <mixed-citation>Bücherl W (1949) Estudos sobre escutigeromorfos Brasileiros. Memórias do Instituto Butantan 21: 9–54.</mixed-citation>
      </ref>
      <ref id="B6">
        <mixed-citation>Castresana J (2000) Selection of conserved blocks from multiple alignments for their use in phylogenetic analysis. Molecular Biology and Evolution 17(4): 540–552. <ext-link xlink:href="10.1093/oxfordjournals.molbev.a026334" ext-link-type="doi">https://doi.org/10.1093/oxfordjournals.molbev.a026334</ext-link></mixed-citation>
      </ref>
      <ref id="B7">
        <mixed-citation>Chamberlin RV (1920) The Myriopoda of the Australian region. Bulletin of the Museum of Comparative Zoology, Harvard College 64: 1–269.</mixed-citation>
      </ref>
      <ref id="B8">
        <mixed-citation>Chamberlin RV (1955) Reports of the Lund University Chile Expedition 1948-49. 18. The <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name> of the Lund University and California Academy of Science Expeditions. Acta Universitatis Lundensis, Nova Series, Avd. 2 51(5): 1–61.</mixed-citation>
      </ref>
      <ref id="B9">
        <mixed-citation>Chamberlin RV (1958) Millipedes and centipedes from Rennell and Guadalcanal Islands. Natural History of Rennell Island, British Solomon Islands 2: 207–212.</mixed-citation>
      </ref>
      <ref id="B10">
        <mixed-citation>Dyachkov YV (2026) On the centipede genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1904 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scutigeridae</tp:taxon-name-part></tp:taxon-name>) of Middle Asia, Kazakhstan, and Asian Russia. Journal of Insect Biodiversity and Systematics 12 (01): 29–39. <ext-link xlink:href="10.48311/jibs.12.01.29" ext-link-type="doi">https://doi.org/10.48311/jibs.12.01.29</ext-link></mixed-citation>
      </ref>
      <ref id="B11">
        <mixed-citation>Edgecombe GD (2011) The <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Myriapoda</tp:taxon-name-part></tp:taxon-name> Vol. 1: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name> – Taxonomic overview: Order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>. In A. Minelli (Ed.), Treatise on Zoology: Anatomy, Taxonomy, Biology (pp. 363–370). Brill.</mixed-citation>
      </ref>
      <ref id="B12">
        <mixed-citation>Edgecombe GD, Barrow L (2007) A new genus of scutigerid centipede (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>) from Western Australia, with new characters for morphological phylogenetics of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>. Zootaxa 1409: 23–50. <ext-link xlink:href="10.5281/zenodo.175540" ext-link-type="doi">https://doi.org/10.5281/zenodo.175540</ext-link></mixed-citation>
      </ref>
      <ref id="B13">
        <mixed-citation>Edgecombe GD, Giribet G (2006) A century later – A total evidence re-evaluation of the phylogeny of scutigeromorph centipedes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Myriapoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>). Invertebrate Systematics 20(5): 503–525. <ext-link xlink:href="10.1071/IS05044" ext-link-type="doi">https://doi.org/10.1071/IS05044</ext-link></mixed-citation>
      </ref>
      <ref id="B14">
        <mixed-citation>Edgecombe GD, Giribet G (2009) Phylogenetics of scutigeromorph centipedes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Myriapoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>) with implications for species delimitation and historical biogeography of the Australian and New Caledonian faunas. Cladistics 25(4): 406–427. <ext-link xlink:href="10.1111/j.1096-0031.2009.00253.x" ext-link-type="doi">https://doi.org/10.1111/j.1096-0031.2009.00253.x</ext-link></mixed-citation>
      </ref>
      <ref id="B15">
        <mixed-citation>Giribet G, Edgecombe GD (2006) Conflict between datasets and phylogeny of centipedes: an analysis based on seven genes and morphology. Proceedings of the Royal Society B: Biological Sciences 273(1586): 531–538. <ext-link xlink:href="10.1098/rspb.2005.3415" ext-link-type="doi">https://doi.org/10.1098/rspb.2005.3415</ext-link></mixed-citation>
      </ref>
      <ref id="B16">
        <mixed-citation>Giribet G, Edgecombe GD (2013) Stable phylogenetic patterns in scutigeromorph centipedes (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Myriapoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>): dating the diversification of an ancient lineage of terrestrial arthropods. Invertebrate Systematics 27: 485–501. <ext-link xlink:href="10.1071/IS13019" ext-link-type="doi">https://doi.org/10.1071/IS13019</ext-link></mixed-citation>
      </ref>
      <ref id="B17">
        <mixed-citation>Guindon S, Dufayard JF, Lefort V, Anisimova M, Hordijk W, Gascuel O (2010) New algorithms and methods to estimate maximum-likelihood phylogenies: assessing the performance of PhyML 3.0. Systematic Biology 59: 307–321. <ext-link xlink:href="10.1093/sysbio/syq010" ext-link-type="doi">https://doi.org/10.1093/sysbio/syq010</ext-link></mixed-citation>
      </ref>
      <ref id="B18">
        <mixed-citation>Hoang DT, Chernomor O, von Haeseler A, Minh BQ, Vinh LS (2018) UFBoot2: Improving the ultrafast bootstrap approximation. Molecular Biology and Evolution 35(2): 518–522. <ext-link xlink:href="10.1093/molbev/msx281" ext-link-type="doi">https://doi.org/10.1093/molbev/msx281</ext-link></mixed-citation>
      </ref>
      <ref id="B19">
        <mixed-citation>Kalyaanamoorthy S, Minh BQ, Wong TKF, von Haeseler A, Jermiin LS (2017) ModelFinder: Fast model selection for accurate phylogenetic estimates. Nature Methods 14(6): 587–589. <ext-link xlink:href="10.1038/nmeth.4285" ext-link-type="doi">https://doi.org/10.1038/nmeth.4285</ext-link></mixed-citation>
      </ref>
      <ref id="B20">
        <mixed-citation>Koch M, Edgecombe GD (2006) Peristomatic structures in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>): A comparative study, with new characters for higher-level systematics. Zoomorphology 125(3): 187–207. <ext-link xlink:href="10.1007/s00435-006-0027-8" ext-link-type="doi">https://doi.org/10.1007/s00435-006-0027-8</ext-link></mixed-citation>
      </ref>
      <ref id="B21">
        <mixed-citation>Katoh K, Standley DM (2013) MAFFT multiple sequence alignment software version 7: Improvements in performance and usability. Molecular Biology and Evolution 30(4): 772–780. <ext-link xlink:href="10.1093/molbev/mst010" ext-link-type="doi">https://doi.org/10.1093/molbev/mst010</ext-link></mixed-citation>
      </ref>
      <ref id="B22">
        <mixed-citation>Katoh K, Standley DM (2014) MAFFT: Iterative refinement and additional methods. Methods in Molecular Biology 1079: 131–146. <ext-link xlink:href="10.1007/978-1-62703-646-7_8" ext-link-type="doi">https://doi.org/10.1007/978-1-62703-646-7_8</ext-link></mixed-citation>
      </ref>
      <ref id="B23">
        <mixed-citation>Katoh K, Rozewicki J, Yamada KD (2019) MAFFT online service: Multiple sequence alignment, interactive sequence choice and visualization. Briefings in Bioinformatics 20(4): 1160–1166. <ext-link xlink:href="10.1093/bib/bbx108" ext-link-type="doi">https://doi.org/10.1093/bib/bbx108</ext-link></mixed-citation>
      </ref>
      <ref id="B24">
        <mixed-citation>Katoh K, Kuma K, Toh H, Miyata T (2005) MAFFT version 5: Improvement in accuracy of multiple sequence alignment. Nucleic Acids Research 33(2): 511–518. <ext-link xlink:href="10.1093/nar/gki198" ext-link-type="doi">https://doi.org/10.1093/nar/gki198</ext-link></mixed-citation>
      </ref>
      <ref id="B25">
        <mixed-citation>Lamarck JB (1801) Système des animaux sans vertèbres. Paris, Deterville, VIII + 432 pp.</mixed-citation>
      </ref>
      <ref id="B26">
        <mixed-citation>Letunic I, Bork P (2021) Interactive Tree of Life (iTOL) v5: An online tool for phylogenetic tree display and annotation. Nucleic Acids Research 49(W1): W293–W296. <ext-link xlink:href="10.1093/nar/gkab301" ext-link-type="doi">https://doi.org/10.1093/nar/gkab301</ext-link></mixed-citation>
      </ref>
      <ref id="B27">
        <mixed-citation>Li Q, Edgecombe GD, Jiang C (2026) A new genus of scutigerid centipede from southern China and its phylogenetic implications (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scutigeridae</tp:taxon-name-part></tp:taxon-name>). Zoologischer Anzeiger 322: 198–211. <ext-link xlink:href="10.1016/j.jcz.2026.04.004" ext-link-type="doi">https://doi.org/10.1016/j.jcz.2026.04.004</ext-link></mixed-citation>
      </ref>
      <ref id="B28">
        <mixed-citation>Manivannan M, Gurung N, Edgecombe GD, Joshi J (2024) A passage through India: The biotic ferry model supports the build-up of Indo-Australian biodiversity of an ancient soil arthropod clade. Journal of Biogeography 51: 2395–2411. <ext-link xlink:href="10.1111/jbi.14994" ext-link-type="doi">https://doi.org/10.1111/jbi.14994</ext-link></mixed-citation>
      </ref>
      <ref id="B29">
        <mixed-citation>Minh BQ, Schmidt HA, Chernomor O, Schrempf D, Woodhams MD, von Haeseler A, Lanfear R (2020) IQ-TREE 2: new models and efficient methods for phylogenetic inference in the genomic era. Molecular Biology and Evolution 37: 1530–1534. <ext-link xlink:href="10.1093/molbev/msaa015" ext-link-type="doi">https://doi.org/10.1093/molbev/msaa015</ext-link></mixed-citation>
      </ref>
      <ref id="B30">
        <mixed-citation>Nguyen LT, Schmidt HA, von Haeseler A, Minh BQ (2015) IQ-TREE: A fast and effective stochastic algorithm for estimating maximum-likelihood phylogenies. Molecular Biology and Evolution 32(1): 268–274. <ext-link xlink:href="10.1093/molbev/msu300" ext-link-type="doi">https://doi.org/10.1093/molbev/msu300</ext-link></mixed-citation>
      </ref>
      <ref id="B31">
        <mixed-citation>Porta AO, Giribet G (2024) A new genus of scutigerid centipede from southern South America with the description of two new species and an updated molecular phylogeny of the myriapod order <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subphylum">Myriapoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>). Invertebrate Systematics 38(4): 1–31. <ext-link xlink:href="10.1071/IS24006" ext-link-type="doi">https://doi.org/10.1071/IS24006</ext-link></mixed-citation>
      </ref>
      <ref id="B32">
        <mixed-citation>Ronquist F, Teslenko M, van der Mark P, Ayres DL, Darling A, Höhna S, Larget B, Liu L, Suchard MA, Huelsenbeck JP (2012) MrBayes 3.2: Efficient Bayesian phylogenetic inference and model choice across a large model space. Systematic Biology 61(3): 539–542. <ext-link xlink:href="10.1093/sysbio/sys029" ext-link-type="doi">https://doi.org/10.1093/sysbio/sys029</ext-link></mixed-citation>
      </ref>
      <ref id="B33">
        <mixed-citation>Sombke A, Rosenberg J, Hilken G, Westermann M, Ernst A (2011) The source of chilopod sensory information: external structure and distribution of antennal sensilla in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Scutigera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">coleoptrata</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>). Journal of Morphology 272: 1376–1387. <ext-link xlink:href="10.1002/jmor.10999" ext-link-type="doi">https://doi.org/10.1002/jmor.10999</ext-link></mixed-citation>
      </ref>
      <ref id="B34">
        <mixed-citation>Trifinopoulos J, Nguyen LT, von Haeseler A, Minh BQ (2016) W-IQ-TREE: a fast online phylogenetic tool for maximum likelihood analysis. Nucleic Acids Research 44: W232–W235. <ext-link xlink:href="10.1093/nar/gkw256" ext-link-type="doi">https://doi.org/10.1093/nar/gkw256</ext-link></mixed-citation>
      </ref>
      <ref id="B35">
        <mixed-citation>Verhoeff KW (1904) Über Gattungen der Spinnenasseln (Scutigeriden). Sitzungs-Berichte der Gesellschaft Naturforschender Freunde zu Berlin 1904: 245–285.</mixed-citation>
      </ref>
      <ref id="B36">
        <mixed-citation>Verhoeff KW (1905) Über Scutigeriden, 5. Zoologischer Anzeiger 29: 73–371.</mixed-citation>
      </ref>
      <ref id="B37">
        <mixed-citation>Verhoeff KW (1925) Results of Dr. E. Mjöberg’s Swedish Scientific Expeditions to Australia 1910–1913. 39. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>. Arkiv för Zoologi 17A(3): 1–62.</mixed-citation>
      </ref>
      <ref id="B38">
        <mixed-citation>Verhoeff KW (1944) Eine neue Scutigeriden-Gattung aus Brasilien. Zoologischer Anzeiger 144: 195–200.</mixed-citation>
      </ref>
      <ref id="B39">
        <mixed-citation>Würmli M (1973) Zur Systematik der Scutigeriden Europas und Kleinasiens (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>). Vorarbeiten zu einer Monographie der <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name>. I. Annalen des Naturhistorischen Museums Wien 77: 399–408.</mixed-citation>
      </ref>
      <ref id="B40">
        <mixed-citation>Würmli M (1974) <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Pesvarus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">pachypus</tp:taxon-name-part></tp:taxon-name></italic> n. gen. n. sp. eine neue Scutigeride (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scutigeridae</tp:taxon-name-part></tp:taxon-name>) aus Australien. Zoologischer Anzeiger 192: 138–146.</mixed-citation>
      </ref>
      <ref id="B41">
        <mixed-citation>Würmli M (1975a) Revision der Hundertfüsser-Gattung <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Thereuonema</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Chilopoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Scutigeridae</tp:taxon-name-part></tp:taxon-name>). Entomologica Germanica 2: 189–196.</mixed-citation>
      </ref>
      <ref id="B42">
        <mixed-citation>Würmli M (1975b) <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Scutigeromorpha</tp:taxon-name-part></tp:taxon-name> von Madagaskar. Die Identität von <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Lassophora</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">madagascariensis</tp:taxon-name-part></tp:taxon-name></italic> Verhoeff, 1905. Bolletino della Società Entomologica Italiana 107(3–5): 70–74.</mixed-citation>
      </ref>
      <ref id="B43">
        <mixed-citation>Zhang D, Gao F, Jakovlić I, Zou H, Zhang J, Li WX, Wang GT (2020) PhyloSuite: An integrated and scalable desktop platform for streamlined molecular sequence data management and evolutionary phylogenetics studies. Molecular Ecology Resources 20(1): 348–355. <ext-link xlink:href="10.1111/1755-0998.13096" ext-link-type="doi">https://doi.org/10.1111/1755-0998.13096</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">10.3897/asp.84.e186815.suppl1</object-id>
        <object-id content-type="arpha">B9BFC8A1-3F11-5544-BCCE-155284CE7DEA</object-id>
        <label>Supplementary Material 1</label>
        <caption>
          <p>Table S1</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p><bold/>: .docx</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation notes</label>
          <p><bold/>: Scutigeromorph and outgroup sampling, MCZ or other accession numbers, country/region of origin and sequenced gene fragments..</p>
        </statement>
        <media xlink:href="arthropod-systematics-84-447-s001.docx" mimetype="application" mime-subtype="vnd.openxmlformats-officedocument.wordprocessingml.document" position="float" orientation="portrait" id="oo_1690402.docx">
          <uri content-type="original_file">https://binary.pensoft.net/file/1690402</uri>
        </media>
        <permissions>
          <license>
            <license-p>This dataset is made available under the Open Database License (<ext-link ext-link-type="uri" xlink:href="http://opendatacommons.org/licenses/odbl/1.0">http://opendatacommons.org/licenses/odbl/1.0</ext-link>). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors"> Li Q, Edgecombe GD, Jiang C (2026)</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
