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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">6</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:DE333E49-E4EE-5693-9BEF-D446E6301C98</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:55528528-5C97-4D79-A718-8B3D153B37CC</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Journal of Hymenoptera Research</journal-title>
        <abbrev-journal-title xml:lang="en">JHR</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="ppub">1070-9428</issn>
      <issn pub-type="epub">1314-2607</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/jhr.91.82490</article-id>
      <article-id pub-id-type="publisher-id">82490</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Formicidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Systematics</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Asia</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>Discovery of a new phragmotic species of the ant genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> Westwood, 1840 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Formicidae</tp:taxon-name-part></tp:taxon-name>) from Cambodia</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Hosoishi</surname>
            <given-names>Shingo</given-names>
          </name>
          <email xlink:type="simple">hosoishi@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-2813-9202</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/investigation/">Investigation</role>
          <role content-type="http://credit.niso.org/contributor-roles/writing-original-draft/">Writing - original draft</role>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Yamane</surname>
            <given-names>Seiki</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>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Sokh</surname>
            <given-names>Heng</given-names>
          </name>
          <xref ref-type="aff" rid="A3">3</xref>
          <role content-type="http://credit.niso.org/contributor-roles/investigation/">Investigation</role>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Institute of Tropical Agriculture, Kyushu University, Motooka 744, Nishi-ku, Fukuoka, 8190395 Japan</addr-line>
        <institution>Kyushu University</institution>
        <addr-line content-type="city">Fukuoka</addr-line>
        <country>Japan</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Haruyama-cho, Kagoshima, Japan</addr-line>
        <institution>Unaffiliated</institution>
        <addr-line content-type="city">Kagoshima</addr-line>
        <country>Japan</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Forestry Administration of Cambodia, 40 Preah Norodom Boulevard, Phnom Penh, Cambodia</addr-line>
        <institution>Forestry Administration of Cambodia</institution>
        <addr-line content-type="city">Phnom Penh</addr-line>
        <country>Cambodia</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Shingo Hosoishi (<email xlink:type="simple">hosoishi@gmail.com</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Francisco Hita Garcia</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2022</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>30</day>
        <month>06</month>
        <year>2022</year>
      </pub-date>
      <volume>91</volume>
      <fpage>357</fpage>
      <lpage>374</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/94B90966-9A51-551E-A953-8D7EACDA7E12">94B90966-9A51-551E-A953-8D7EACDA7E12</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/B460C9E8-7259-4063-BF09-0338DA6C8119">B460C9E8-7259-4063-BF09-0338DA6C8119</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/6789868">6789868</uri>
      <history>
        <date date-type="received">
          <day>19</day>
          <month>02</month>
          <year>2022</year>
        </date>
        <date date-type="accepted">
          <day>15</day>
          <month>05</month>
          <year>2022</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Shingo Hosoishi, Seiki Yamane, Heng Sokh</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">http://zoobank.org/B460C9E8-7259-4063-BF09-0338DA6C8119</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>Several phragmotic species in the ant genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> Westwood, 1840 with a cephalic shield are known from the Old World, but species with an anteriorly truncated head seem to be unknown until now. A new species, resembling the phragmotic workers of the ant genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobopsis">Colobopsis</tp:taxon-name-part></tp:taxon-name></italic>, is here described as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> Hosoishi &amp; Yamane, <bold>sp. nov.</bold>, based on major and minor worker found in Cambodia. This new species is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> (Xu) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="obtusidenta">obtusidenta</tp:taxon-name-part></tp:taxon-name></italic> (Xu) in some important characters, but distinguished from the latter two by distinct metanotum and deep groove in front of metanotal disc in the major worker, and longer head and deep metanotal groove in the minor worker. This is the first confirmed record of a truncated phragmotic head in the major worker of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic>. We propose the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> species group to include all the three species, present a diagnosis of this group, and provide a key to species based on the major and minor worker castes.</p>
      </abstract>
      <kwd-group>
        <label>Keywords</label>
        <kwd>Ants</kwd>
        <kwd>new species</kwd>
        <kwd>phragmosis</kwd>
        <kwd>Southeast Asia</kwd>
        <kwd>taxonomy</kwd>
        <kwd>truncated head</kwd>
      </kwd-group>
    </article-meta>
    <notes>
      <sec sec-type="Citation" id="SECID0EWG">
        <title>Citation</title>
        <p>Hosoishi S, Yamane Sk, Sokh H (2022) Discovery of a new phragmotic species of the ant genus <italic>Carebara</italic> Westwood, 1840 (Hymenoptera, Formicidae) from Cambodia. Journal of Hymenoptera Research 91: 357–374. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.3897/jhr.91.82490">https://doi.org/10.3897/jhr.91.82490</ext-link></p>
      </sec>
    </notes>
  </front>
  <body>
    <sec sec-type="Introduction" id="SECID0EXH">
      <title>Introduction</title>
      <p>The ant genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> Westwood, 1840 belongs to the tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crematogastrini</tp:taxon-name-part></tp:taxon-name> of the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part></tp:taxon-name> (<xref ref-type="bibr" rid="B27">Ward et al. 2015</xref>). The genus is distributed worldwide, mainly in tropical and subtropical regions, and contains more than 200 described taxa (<xref ref-type="bibr" rid="B4">Bolton 2022</xref>). The taxonomy of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> in the Afrotropical and Malagasy regions was remarkably improved in recent years (<xref ref-type="bibr" rid="B10">Fischer et al. 2014</xref>, <xref ref-type="bibr" rid="B11">2015</xref>; <xref ref-type="bibr" rid="B2">Azorsa and Fisher 2018</xref>). In Asia, several taxonomic studies have been conducted for species of Japan (<xref ref-type="bibr" rid="B24">Terayama et al. 2014</xref>), China (<xref ref-type="bibr" rid="B17">Li and Tang 1986</xref>; <xref ref-type="bibr" rid="B31">Wu and Wang 1995</xref>; <xref ref-type="bibr" rid="B35">Zhou and Zheng 1997</xref>; <xref ref-type="bibr" rid="B33">Xu 2003</xref>; <xref ref-type="bibr" rid="B34">Zhou et al. 2006</xref>), Taiwan (<xref ref-type="bibr" rid="B23">Terayama et al. 2012</xref>), Thailand (<xref ref-type="bibr" rid="B16">Jaitrong et al. 2021</xref>), and India (<xref ref-type="bibr" rid="B3">Bharti and Kumar 2013</xref>). In contrast, little attention has been paid to the species of Southeast Asia except for Thailand, one of the species-rich areas for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic>, especially for species previously included in the now synonymized <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Oligomyrmex">Oligomyrmex</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidologeton">Pheidologeton</tp:taxon-name-part></tp:taxon-name></italic>. Species-level identifications are challenging due to a lack of comprehensive revisions of the genus in all the Old World except Madagascar (<xref ref-type="bibr" rid="B11">Fischer et al. 2015</xref>; <xref ref-type="bibr" rid="B2">Azorsa and Fisher 2018</xref>), although species with unique morphological characters have been described in smaller taxonomic treatments (e.g., <xref ref-type="bibr" rid="B1">Akbar and Bharti 2017</xref>; <xref ref-type="bibr" rid="B16">Jaitrong et al. 2021</xref>).</p>
      <p>Among the unique morphological characters in ants, one of the most representative is ‘phragmosis’ (<xref ref-type="bibr" rid="B29">Wheeler 1927</xref>). Phragmosis is a method in which an animal defends itself in its burrow or nest by using its own modified body structure to plug the entrances. In ants (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Formicidae</tp:taxon-name-part></tp:taxon-name>), phragmosis is widely known from several genera, i.e. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blepharidatta">Blepharidatta</tp:taxon-name-part></tp:taxon-name></italic> Wheeler, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobopsis">Colobopsis</tp:taxon-name-part></tp:taxon-name></italic> Mayr, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cephalotes">Cephalotes</tp:taxon-name-part></tp:taxon-name></italic> Latreille, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobostruma">Colobostruma</tp:taxon-name-part></tp:taxon-name></italic> Wheeler, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Crematogaster">Crematogaster</tp:taxon-name-part></tp:taxon-name></italic> Lund, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part></tp:taxon-name></italic> Westwood, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tetraponera">Tetraponera</tp:taxon-name-part></tp:taxon-name></italic> Smith, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> Westwood (<xref ref-type="bibr" rid="B15">Hölldobler and Wilson 1990</xref>; <xref ref-type="bibr" rid="B11">Fischer et al. 2015</xref>). <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref> revised the African phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species with a cephalic shield in a special major worker subcaste. Their examinations suggested that the phragmosis in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> is not always precisely displayed by major workers, but sometimes by ergatoid queens. Other phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species are known from South Asia (<xref ref-type="bibr" rid="B12">Forel 1902</xref>, <xref ref-type="bibr" rid="B13">1913</xref>; <xref ref-type="bibr" rid="B21">Sheela and Narendran 1997</xref>). <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref> suggested that phragmosis in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> evolved independently in the Old World. In these cases, phragmotic workers have the same type of concave cephalic shield on the dorsal surface of the head. The cephalic shield is formed by the depressed dorsal surface of the head, clypeus and mandibles in the major worker subcastes of those species. Whereas an anteriorly truncated head, such as in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobopsis">Colobopsis</tp:taxon-name-part></tp:taxon-name></italic>, has not been reported for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> until now.</p>
      <p>Recently, we examined <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> specimens of a phragmotic species collected from a dry evergreen forest in Cambodia. The species is considered to be distinct and new to science in that its majors have an anteriorly truncated head, which clearly distinguishes it from other Asian species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic>. In this paper, we describe this new species based on the major and minor worker subcastes. It is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> (Xu) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="obtusidenta">obtusidenta</tp:taxon-name-part></tp:taxon-name></italic> (Xu), which have been described from Yunnan Province, China (<xref ref-type="bibr" rid="B33">Xu 2003</xref>). In this paper the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic>-group is established, and its diagnosis and a key to species are given based on the major and minor worker castes.</p>
    </sec>
    <sec sec-type="methods" id="SECID0ERJAC">
      <title>Methods</title>
      <p>Most observations were made on a Leica M205C stereomicroscope. Images were taken using a Canon EOS 50D with a Canon MP-E 65 mm 1–5 × Macro lens and a Leica DFC 450 digital imaging system, then processed using Combine ZM (<xref ref-type="bibr" rid="B14">Hadley 2010</xref>).</p>
      <p>Specimens used in this study were collected during expeditions in Cambodia under the Memorandum of Understanding (<bold><abbrev xlink:title="Memorandum of Understanding" id="ABBRID0E5JAC">MOU</abbrev></bold>) between Japan and Cambodia, Kyushu University, Japan and the Forestry Administration, Cambodia, on cooperation concerning biological resources and information. The materials are shared between Cambodia and Japan, but the present materials are deposited in Japan, on indefinite loan from Cambodia. Type specimens were examined and/or deposited in the Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan (<bold><abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0EDKAC">KUEC</abbrev></bold>).</p>
      <p>Measurements were made under a Leica M205C stereomicroscope using micrometers. All measurements are expressed in millimeters, recorded to the second decimal place. Measurements and morphological terminology follow <xref ref-type="bibr" rid="B10">Fischer et al. (2014)</xref> and <xref ref-type="bibr" rid="B2">Azorsa and Fisher (2018)</xref>. Abbreviations of measurements and indices are as follows:</p>
      <p><bold><abbrev xlink:title="head length" id="ABBRID0EVKAC">HL</abbrev></bold> head length: maximum length of head in full-face view between lines drawn across anterior margin of clypeus and lines drawn across the posterolateral corners of head;</p>
      <p><bold><abbrev xlink:title="head width" id="ABBRID0E3KAC">HW</abbrev></bold> head width: width of head directly behind the eyes measured in full-face view;</p>
      <p><bold><abbrev xlink:title="scape length" id="ABBRID0EDLAC">SL</abbrev></bold> scape length: maximum scape length excluding basal condyle and neck;</p>
      <p><bold><abbrev xlink:title="mandible length" id="ABBRID0EKLAC">ML</abbrev></bold> mandible length: the straight-line length of the mandible at full closure from the mandibular apex to the clypeal margin;</p>
      <p><bold><abbrev xlink:title="eye length" id="ABBRID0ERLAC">EL</abbrev></bold> eye length: maximum diameter of compound eye measured in oblique profile view;</p>
      <p><bold><abbrev xlink:title="eye to mandible" id="ABBRID0EYLAC">EM</abbrev></bold> eye to mandible: distance from base of compound eye to the mandibular insertion, measured in profile view;</p>
      <p><bold><abbrev xlink:title="head depth" id="ABBRID0E6LAC">HD</abbrev></bold> head depth: maximum depth of head in profile view measured perpendicular to full-face view plane;</p>
      <p><bold><abbrev xlink:title="Weber’s length" id="ABBRID0EGMAC">WL</abbrev></bold> Weber’s length: diagonal length of mesosoma in profile view from the postero-ventral margin of the propodeal lobe to the anterior-most point of the pronotal slope, excluding the neck;</p>
      <p><bold><abbrev xlink:title="propodeal spine length" id="ABBRID0ENMAC">PSL</abbrev></bold> propodeal spine length: length of propodeal spine or posterodorsal corner of propodeum, measured from center of the propodeal spiracle to the tip of the spine of the posterodorsal corner of propodeum;</p>
      <p><bold><abbrev xlink:title="pronotal width" id="ABBRID0EUMAC">PW</abbrev></bold> pronotal width: maximum width of pronotum measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="femur length" id="ABBRID0E2MAC">MFL</abbrev></bold> femur length: length of the profemur measured along its long axis in posterior view;</p>
      <p><bold><abbrev xlink:title="femur width" id="ABBRID0ECNAC">MFW</abbrev></bold> femur width: maximum width of the hind femur;</p>
      <p><bold><abbrev xlink:title="tibia length" id="ABBRID0EJNAC">MTL</abbrev></bold> tibia length: maximum length of hind tibia;</p>
      <p><bold><abbrev xlink:title="petiole length" id="ABBRID0EQNAC">PTL</abbrev></bold> petiole length: maximum length of petiole, measured in profile view;</p>
      <p><bold><abbrev xlink:title="petiolar node height" id="ABBRID0EXNAC">PTH</abbrev></bold> petiolar node height: maximum height of petiolar node measured in profile view from the highest (median) point of the node to the ventral outline;</p>
      <p><bold><abbrev xlink:title="petiolar node length" id="ABBRID0E5NAC">PNL</abbrev></bold> petiolar node length: maximum length of petiolar node measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="petiolar node width" id="ABBRID0EFOAC">PTW</abbrev></bold> petiolar node width: maximum width of dorsal face of petiolar node measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="propodeal spiracle diameter" id="ABBRID0EMOAC">PSD</abbrev></bold> propodeal spiracle diameter: maximum length of propodeal spiracle in profile view;</p>
      <p><bold><abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0ETOAC">DPSD</abbrev></bold> distance from propodeal spiracle to declivity: minimum distance from posterior border of propodeal spiracle to declivity of propodeum;</p>
      <p><bold><abbrev xlink:title="postpetiole length" id="ABBRID0E1OAC">PPL</abbrev></bold> postpetiole length: maximum length of postpetiole measured in profile view;</p>
      <p><bold><abbrev xlink:title="postpetiole node length" id="ABBRID0EBPAC">PPNL</abbrev></bold> postpetiole node length: maximum length of postpetiole measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="postpetiole height" id="ABBRID0EIPAC">PPH</abbrev></bold> postpetiole height: maximum height of the postpetiole measured in profile view from the highest (median) point of the node to the ventral outline. The measuring line is placed at an orthogonal angle to the ventral outline of the node;</p>
      <p><bold><abbrev xlink:title="postpetiole width" id="ABBRID0EPPAC">PPW</abbrev></bold> postpetiole width: maximum width of postpetiole measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="gaster length" id="ABBRID0EWPAC">GL</abbrev></bold> gaster length: maximum length of the gaster measured in profile view;</p>
      <p><bold><abbrev xlink:title="gaster width" id="ABBRID0E4PAC">GW</abbrev></bold> gaster width: maximum width of the gaster measured in dorsal view;</p>
      <p><bold><abbrev xlink:title="cephalic index" id="ABBRID0EFAAE">CI</abbrev></bold> cephalic index: 100*<abbrev xlink:title="head width" id="ABBRID0EJAAE">HW</abbrev>/<abbrev xlink:title="head length" id="ABBRID0ENAAE">HL</abbrev>;</p>
      <p><bold><abbrev xlink:title="mandibular index" id="ABBRID0EUAAE">MI</abbrev></bold> mandibular index: 100*<abbrev xlink:title="mandible length" id="ABBRID0EYAAE">ML</abbrev>/<abbrev xlink:title="head length" id="ABBRID0E3AAE">HL</abbrev>;</p>
      <p><bold><abbrev xlink:title="scape index" id="ABBRID0EDBAE">SI</abbrev></bold> scape index: 100*<abbrev xlink:title="scape length" id="ABBRID0EHBAE">SL</abbrev>/<abbrev xlink:title="head length" id="ABBRID0ELBAE">HL</abbrev>;</p>
      <p><bold><abbrev xlink:title="metafemur length index" id="ABBRID0ESBAE">MLI</abbrev></bold> metafemur length index: 100*FL/<abbrev xlink:title="head length" id="ABBRID0EWBAE">HL</abbrev>;</p>
      <p><bold><abbrev xlink:title="postpetiole length index" id="ABBRID0E4BAE">PPLI</abbrev></bold> postpetiole length index: 100*<abbrev xlink:title="postpetiole length" id="ABBRID0EBCAE">PPL</abbrev>/<abbrev xlink:title="petiole length" id="ABBRID0EFCAE">PTL</abbrev>;</p>
      <p><bold><abbrev xlink:title="postpetiole width index" id="ABBRID0EMCAE">PPI</abbrev></bold> postpetiole width index: 100*<abbrev xlink:title="postpetiole width" id="ABBRID0EQCAE">PPW</abbrev>/<abbrev xlink:title="petiolar node width" id="ABBRID0EUCAE">PTW</abbrev>;</p>
      <p><bold><abbrev xlink:title="propodeal spine index" id="ABBRID0E2CAE">PSI</abbrev></bold> propodeal spine index: 100*<abbrev xlink:title="propodeal spine length" id="ABBRID0E6CAE">PSL</abbrev>/<abbrev xlink:title="head width" id="ABBRID0EDDAE">HW</abbrev>.</p>
    </sec>
    <sec sec-type="Results" id="SECID0EHDAE">
      <title>Results</title>
      <sec sec-type="The Carebara acutispina-group" id="SECID0ELDAE">
        <title>The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic>-group</title>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> (Xu, 2003)</p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> Hosoishi &amp; Yamane, sp. nov.</p>
        <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="obtusidenta">obtusidenta</tp:taxon-name-part></tp:taxon-name></italic> (Xu, 2003)</p>
        <p><bold>Diagnosis.</bold> The <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> group can be distinguished from other Asian <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species by the following character conditions: (1) mandible with five teeth, but six teeth in the phragmotic major worker in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic>; (2) 9-segmented antennae; (3) posterolateral corner of head with a horn in major worker; (4) promesonotum convex in major worker; (5) developed propodeal spines; (6) propodeal declivity with lateral lamellae.</p>
        <p><bold>Notes.</bold> The members of this group are similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="wheeleri">wheeleri</tp:taxon-name-part></tp:taxon-name></italic> (Ettershank) in having features (2), (3) and (5), but can be distinguished from the latter by having the aforementioned feature (1). The species group is known from southern China and Cambodia, but has not been recorded from the neighboring regions (e.g., Vietnam, Thailand).</p>
      </sec>
      <sec sec-type="Key to the Carebara acutispina-group species based on the major worker" id="SECID0ESGAE">
        <title>Key to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic>-group species based on the major worker</title>
        <p>The following key is partly based on <xref ref-type="bibr" rid="B33">Xu (2003)</xref>.</p>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0EVIAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Head phragmotic, distinctly truncate anteriorly, in full-face view with deeply concave posterior margin. Promesonotum strongly convex dorsally; metanotal disc present, groove in front of the disc deep; propodeal spine distinct; subpetiolar process obsolete</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Head normal, not truncated anteriorly, in full-face view with shallowly emarginated posterior margin</td>
                <td rowspan="1" colspan="1">
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Vertex without transverse rugulae. Promesonotum moderately convex dorsally. Metanotal groove shallow. Propodeal spine distinct. Subpetiolar process obsolete</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> (Xu)</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Vertex with transverse rugulae. Promesonotum highly convex dorsally. Metanotal groove deep. Propodeal spine rudimentary. Subpetiolar process more developed</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="obtusidenta">obtusidenta</tp:taxon-name-part></tp:taxon-name></italic> (Xu)</bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
      <sec sec-type="Key to the Carebara acutispina-group species based on the minor worker" id="SECID0EIKAE">
        <title>Key to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic>-group species based on the minor worker</title>
        <p>The following key is modified from <xref ref-type="bibr" rid="B33">Xu (2003)</xref>.</p>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0EDMAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Head relatively longer (<abbrev xlink:title="cephalic index" id="ABBRID0EOLAE">CI</abbrev> 81–84). Metanotal groove deep, U-shaped in section</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Head relatively wider (<abbrev xlink:title="cephalic index" id="ABBRID0EMMAE">CI</abbrev> 85–93). Metanotal groove shallow, V-shaped in section</td>
                <td rowspan="1" colspan="1">
                  <bold>2</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Body smaller (<abbrev xlink:title="head length" id="ABBRID0E5MAE">HL</abbrev> 0.30–0.33, <abbrev xlink:title="head width" id="ABBRID0ECNAE">HW</abbrev> 0.28, <abbrev xlink:title="scape length" id="ABBRID0EGNAE">SL</abbrev> 0.18–0.19, <abbrev xlink:title="pronotal width" id="ABBRID0EKNAE">PW</abbrev> 0.18–0.20). Dorsum of propodeum weakly depressed</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> (Xu)</bold>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Body larger (<abbrev xlink:title="head length" id="ABBRID0EIOAE">HL</abbrev> 0.38–0.40, <abbrev xlink:title="head width" id="ABBRID0EMOAE">HW</abbrev> 0.33–0.34, <abbrev xlink:title="scape length" id="ABBRID0EQOAE">SL</abbrev> 0.23–0.25, <abbrev xlink:title="pronotal width" id="ABBRID0EUOAE">PW</abbrev> 0.23–0.24). Dorsum of propodeum evenly convex</td>
                <td rowspan="1" colspan="1">
                  <bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="obtusidenta">obtusidenta</tp:taxon-name-part></tp:taxon-name></italic> (Xu)</bold>
                </td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
        <tp:taxon-treatment>
          <tp:treatment-meta>
            <kwd-group>
              <label>Taxon classification</label>
              <kwd>
                <named-content content-type="kingdom" xlink:type="simple">Animalia</named-content>
              </kwd>
              <kwd>
                <named-content content-type="order" xlink:type="simple">Hymenoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Formicidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">4AB0F3C2-8703-5FF5-9F22-46B14E092EA3</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/62D5C172-3C04-477D-B7DC-6395F018250C</object-id>
            </tp:taxon-name>
            <tp:taxon-authority>Hosoishi &amp; Yamane</tp:taxon-authority>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1</xref>
            <xref ref-type="fig" rid="F2">, 2</xref>
            <xref ref-type="fig" rid="F3">, 3</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="type material" id="SECID0E6QAE">
            <title>Type material.</title>
            <p><abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0EFRAE">KUEC</abbrev>; <bold><italic>Holotype</italic></bold> (major worker); Cambodia: Siem Reap Province, Local Forest Administration plot, <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[104.043889,13.729722]}" id="NCID0EQRAE">13°43'47"N, 104°02'38"E</named-content></named-content>, 13. XII. 2012, S. Hosoishi leg., SH12-Cam-135 (collection code KUECANT031). – <abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0EVRAE">KUEC</abbrev>; <bold><italic>Paratypes</italic></bold>: 2 major workers (collection code KUECANT032, KUECANT033) (<abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0E4RAE">KUEC</abbrev>), 4 minor workers (collection code KUECANT034, KUECANT035, KUECANT036, KUECANT037) (<abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0ECSAE">KUEC</abbrev>), same data as holotype.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type material" id="SECID0EHSAE">
            <title>Non-type material examined.</title>
            <p>Three minor workers (collection code KUECANT038, KUECANT039, KUECANT040) (<abbrev content-type="institution" xlink:title="Institute of Tropical Agriculture, Kyushu University, Fukuoka, Japan" id="ABBRID0ENSAE">KUEC</abbrev>), same data as holotype.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0ESSAE">
            <title>Diagnosis of worker.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> can be distinguished from the other species in the genus by the following characters: truncated head, posterolateral corner of head with a horn, 9-segmented antennae, distinct metanotum and deep metanotal groove, propodeal declivity with lamella laterally in the major worker subcaste; longer head, 9-segmented antennae, smooth surface of promesonotum, deep metanotal groove, propodeum armed with a long acute tooth, propodeal declivity with lamella and petiolar venter with lamella along its length in the minor worker subcaste.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EDTAE">
            <title>Measurements and indices.</title>
            <p>Holotype major worker: <abbrev xlink:title="head length" id="ABBRID0EJTAE">HL</abbrev> 0.84; <abbrev xlink:title="head width" id="ABBRID0ENTAE">HW</abbrev> 0.63; <abbrev xlink:title="scape length" id="ABBRID0ERTAE">SL</abbrev> 0.29; <abbrev xlink:title="mandible length" id="ABBRID0EVTAE">ML</abbrev> 0.22; <abbrev xlink:title="eye length" id="ABBRID0EZTAE">EL</abbrev> 0.05; <abbrev xlink:title="eye to mandible" id="ABBRID0E4TAE">EM</abbrev> 0.22; <abbrev xlink:title="head depth" id="ABBRID0EBUAE">HD</abbrev> 0.53; <abbrev xlink:title="Weber’s length" id="ABBRID0EFUAE">WL</abbrev> 0.68; <abbrev xlink:title="propodeal spine length" id="ABBRID0EJUAE">PSL</abbrev> 0.13; <abbrev xlink:title="pronotal width" id="ABBRID0ENUAE">PW</abbrev> 0.35; <abbrev xlink:title="femur length" id="ABBRID0ERUAE">MFL</abbrev> 0.41; <abbrev xlink:title="femur width" id="ABBRID0EVUAE">MFW</abbrev> 0.11; <abbrev xlink:title="tibia length" id="ABBRID0EZUAE">MTL</abbrev> 0.33; <abbrev xlink:title="petiole length" id="ABBRID0E4UAE">PTL</abbrev> 0.26; <abbrev xlink:title="petiolar node height" id="ABBRID0EBVAE">PTH</abbrev> 0.18; <abbrev xlink:title="petiolar node length" id="ABBRID0EFVAE">PNL</abbrev> 0.10; <abbrev xlink:title="petiolar node width" id="ABBRID0EJVAE">PTW</abbrev> 0.18; <abbrev xlink:title="propodeal spiracle diameter" id="ABBRID0ENVAE">PSD</abbrev> 0.03; <abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0ERVAE">DPSD</abbrev> 0.06; <abbrev xlink:title="postpetiole length" id="ABBRID0EVVAE">PPL</abbrev> 0.15; <abbrev xlink:title="postpetiole node length" id="ABBRID0EZVAE">PPNL</abbrev> 0.10; <abbrev xlink:title="postpetiole height" id="ABBRID0E4VAE">PPH</abbrev> 0.14; <abbrev xlink:title="postpetiole width" id="ABBRID0EBWAE">PPW</abbrev> 0.19; <abbrev xlink:title="gaster length" id="ABBRID0EFWAE">GL</abbrev> 1.06; <abbrev xlink:title="gaster width" id="ABBRID0EJWAE">GW</abbrev> 0.69; <abbrev xlink:title="cephalic index" id="ABBRID0ENWAE">CI</abbrev> 75; <abbrev xlink:title="mandibular index" id="ABBRID0ERWAE">MI</abbrev> 26; <abbrev xlink:title="scape index" id="ABBRID0EVWAE">SI</abbrev> 34; <abbrev xlink:title="metafemur length index" id="ABBRID0EZWAE">MLI</abbrev> 49; <abbrev xlink:title="postpetiole length index" id="ABBRID0E4WAE">PPLI</abbrev> 55; <abbrev xlink:title="postpetiole width index" id="ABBRID0EBXAE">PPI</abbrev> 106; <abbrev xlink:title="propodeal spine index" id="ABBRID0EFXAE">PSI</abbrev> 21. Paratype major workers (n = 2): <abbrev xlink:title="head length" id="ABBRID0EJXAE">HL</abbrev> 0.82–0.86; <abbrev xlink:title="head width" id="ABBRID0ENXAE">HW</abbrev> 0.63–0.64; <abbrev xlink:title="scape length" id="ABBRID0ERXAE">SL</abbrev> 0.28–0.30; <abbrev xlink:title="mandible length" id="ABBRID0EVXAE">ML</abbrev> 0.20–0.21; <abbrev xlink:title="eye length" id="ABBRID0EZXAE">EL</abbrev> 0.04–0.05; <abbrev xlink:title="eye to mandible" id="ABBRID0E4XAE">EM</abbrev> 0.22–0.23; <abbrev xlink:title="head depth" id="ABBRID0EBYAE">HD</abbrev> 0.53–0.55; <abbrev xlink:title="Weber’s length" id="ABBRID0EFYAE">WL</abbrev> 0.71–0.72; <abbrev xlink:title="propodeal spine length" id="ABBRID0EJYAE">PSL</abbrev> 0.13–0.14; <abbrev xlink:title="pronotal width" id="ABBRID0ENYAE">PW</abbrev> 0.36–0.38; <abbrev xlink:title="femur length" id="ABBRID0ERYAE">MFL</abbrev> 0.41–0.42; <abbrev xlink:title="femur width" id="ABBRID0EVYAE">MFW</abbrev> 0.11–0.12; <abbrev xlink:title="tibia length" id="ABBRID0EZYAE">MTL</abbrev> 0.33–0.35; <abbrev xlink:title="petiole length" id="ABBRID0E4YAE">PTL</abbrev> 0.26–0.28; <abbrev xlink:title="petiolar node height" id="ABBRID0EBZAE">PTH</abbrev> 0.18–0.19; <abbrev xlink:title="petiolar node length" id="ABBRID0EFZAE">PNL</abbrev> 0.09–0.10; <abbrev xlink:title="petiolar node width" id="ABBRID0EJZAE">PTW</abbrev> 0.18–0.19; <abbrev xlink:title="propodeal spiracle diameter" id="ABBRID0ENZAE">PSD</abbrev> 0.03–0.04; <abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0ERZAE">DPSD</abbrev> 0.06–0.07; <abbrev xlink:title="postpetiole length" id="ABBRID0EVZAE">PPL</abbrev> 0.14–0.15; <abbrev xlink:title="postpetiole node length" id="ABBRID0EZZAE">PPNL</abbrev> 0.10–0.12; <abbrev xlink:title="postpetiole height" id="ABBRID0E4ZAE">PPH</abbrev> 0.14–0.15; <abbrev xlink:title="postpetiole width" id="ABBRID0EB1AE">PPW</abbrev> 0.19–0.21; <abbrev xlink:title="gaster length" id="ABBRID0EF1AE">GL</abbrev> 1.00–1.39; <abbrev xlink:title="gaster width" id="ABBRID0EJ1AE">GW</abbrev> 0.48–0.80; <abbrev xlink:title="cephalic index" id="ABBRID0EN1AE">CI</abbrev> 73–76; <abbrev xlink:title="mandibular index" id="ABBRID0ER1AE">MI</abbrev> 23–25; <abbrev xlink:title="scape index" id="ABBRID0EV1AE">SI</abbrev> 33–35; <abbrev xlink:title="metafemur length index" id="ABBRID0EZ1AE">MLI</abbrev> 48–50; <abbrev xlink:title="postpetiole length index" id="ABBRID0E41AE">PPLI</abbrev> 51–56; <abbrev xlink:title="postpetiole width index" id="ABBRID0EB2AE">PPI</abbrev> 106–118; <abbrev xlink:title="propodeal spine index" id="ABBRID0EF2AE">PSI</abbrev> 20–23. Paratype minor workers (n = 4): <abbrev xlink:title="head length" id="ABBRID0EJ2AE">HL</abbrev> 0.36–0.38; <abbrev xlink:title="head width" id="ABBRID0EN2AE">HW</abbrev> 0.30–0.31; <abbrev xlink:title="scape length" id="ABBRID0ER2AE">SL</abbrev> 0.23–0.24; <abbrev xlink:title="mandible length" id="ABBRID0EV2AE">ML</abbrev> 0.08–0.10; <abbrev xlink:title="eye length" id="ABBRID0EZ2AE">EL</abbrev> 0.01; <abbrev xlink:title="eye to mandible" id="ABBRID0E42AE">EM</abbrev> 0.09–0.10; <abbrev xlink:title="head depth" id="ABBRID0EB3AE">HD</abbrev> 0.21–0.22: <abbrev xlink:title="Weber’s length" id="ABBRID0EF3AE">WL</abbrev> 0.36–0.39; <abbrev xlink:title="propodeal spine length" id="ABBRID0EJ3AE">PSL</abbrev> 0.07–0.09; <abbrev xlink:title="pronotal width" id="ABBRID0EN3AE">PW</abbrev> 0.17–0.18; <abbrev xlink:title="femur length" id="ABBRID0ER3AE">MFL</abbrev> 0.23–0.24; <abbrev xlink:title="femur width" id="ABBRID0EV3AE">MFW</abbrev> 0.05–0.06; <abbrev xlink:title="tibia length" id="ABBRID0EZ3AE">MTL</abbrev> 0.18–0.19; <abbrev xlink:title="petiole length" id="ABBRID0E43AE">PTL</abbrev> 0.15–0.16; <abbrev xlink:title="petiolar node height" id="ABBRID0EB4AE">PTH</abbrev> 0.10–0.11; <abbrev xlink:title="petiolar node length" id="ABBRID0EF4AE">PNL</abbrev> 0.06–0.07; <abbrev xlink:title="petiolar node width" id="ABBRID0EJ4AE">PTW</abbrev> 0.06–0.07; <abbrev xlink:title="propodeal spiracle diameter" id="ABBRID0EN4AE">PSD</abbrev> 0.01–0.02; <abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0ER4AE">DPSD</abbrev> 0.01–0.02; <abbrev xlink:title="postpetiole length" id="ABBRID0EV4AE">PPL</abbrev> 0.06–0.07; <abbrev xlink:title="postpetiole node length" id="ABBRID0EZ4AE">PPNL</abbrev> 0.06; <abbrev xlink:title="postpetiole height" id="ABBRID0E44AE">PPH</abbrev> 0.06–0.07; <abbrev xlink:title="postpetiole width" id="ABBRID0EB5AE">PPW</abbrev> 0.08–0.09; <abbrev xlink:title="gaster length" id="ABBRID0EF5AE">GL</abbrev> 0.31–0.34; <abbrev xlink:title="gaster width" id="ABBRID0EJ5AE">GW</abbrev> 0.22–0.26; <abbrev xlink:title="cephalic index" id="ABBRID0EN5AE">CI</abbrev> 81–84; <abbrev xlink:title="mandibular index" id="ABBRID0ER5AE">MI</abbrev> 22–26; <abbrev xlink:title="scape index" id="ABBRID0EV5AE">SI</abbrev> 61–65; <abbrev xlink:title="metafemur length index" id="ABBRID0EZ5AE">MLI</abbrev> 63–67; <abbrev xlink:title="postpetiole length index" id="ABBRID0E45AE">PPLI</abbrev> 41–46; <abbrev xlink:title="postpetiole width index" id="ABBRID0EB6AE">PPI</abbrev> 118–139; <abbrev xlink:title="propodeal spine index" id="ABBRID0EF6AE">PSI</abbrev> 26–28.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/jhr.91.82490.figure1</object-id>
              <object-id content-type="arpha">CEB1826F-55A8-5174-9715-A43D01D96963</object-id>
              <label>Figure 1.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., holotype major worker (KUECANT031). Body in profile view.</p>
              </caption>
              <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707300.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/707300</uri>
              </graphic>
            </fig>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/jhr.91.82490.figure2</object-id>
              <object-id content-type="arpha">6E3E1295-04FE-5687-99C7-82605E84EE33</object-id>
              <label>Figure 2.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., holotype major worker (KUECANT031) <bold>A</bold> body in dorsal view <bold>B</bold> head in full-face view <bold>C</bold> close-up of cephalic shield.</p>
              </caption>
              <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707301.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/707301</uri>
              </graphic>
            </fig>
            <fig id="F3" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/jhr.91.82490.figure3</object-id>
              <object-id content-type="arpha">B877B417-988A-58B8-9EFE-D04F24197C28</object-id>
              <label>Figure 3.</label>
              <caption>
                <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., paratype minor worker (KUECANT034) <bold>A</bold> body in profile view <bold>B</bold> head in full-face view <bold>C</bold> body in dorsal view.</p>
              </caption>
              <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707302.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/707302</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0E5BAG">
            <title>Description of major worker.</title>
            <p>Head, in full-face view, nearly rectangular, long, about 1.3 times as long as wide (<abbrev xlink:title="cephalic index" id="ABBRID0EECAG">CI</abbrev> 73–76), phragmotic with truncated anterior surface of head; posterior margin medially concave; posterolateral corner rounded; lateral margin straight to weakly convex. Posterolateral corner of head with a distinct, acute tooth resembling a horn. Supraclypeal area with deep transverse concavity located just behind posterior margin of clypeus. Clypeus entirely involved in phragmotic shield, margined throughout with continuous high rim; anterior margin medially and laterally very shallowly emarginate. Head, in profile view, gradually narrowed posteriad, with straight anterior margin; dorsal and ventral margins shallowly convex; anterior portion (phragmotic shield) separated from posterior portion by right-angled bent just in front of antennal socket. Ocelli absent. Compound eyes small, consisting of five to six ommatidia (<abbrev xlink:title="eye length" id="ABBRID0EICAG">EL</abbrev> 0.05). Mandibles in frontal view roughly triangular, taking part in round phragmotic shield; masticatory margin straight with six low and blunt teeth; basal margin shallowly convex; outer margin evenly curved. Antenna with nine segments; scape short (<abbrev xlink:title="head length" id="ABBRID0EMCAG">HL</abbrev> 0.82–0.86, <abbrev xlink:title="scape length" id="ABBRID0EQCAG">SL</abbrev> 0.28–0.30, <abbrev xlink:title="scape index" id="ABBRID0EUCAG">SI</abbrev> 33–35) not surpassing cephalic midlength, basally narrow and widening apicad; third to seventh antennal segments short, globular; apical two segments forming distinct club.</p>
            <p>With mesosoma in dorsal view, pronotum globular, large, occupying most of mesosoma; mesonotum small, transverse, nearly as wide as propodeum; promesonotal suture faint; metanotum forming narrow, elevated band; dorsal face of propodeum wider than long; not defined from posterior declivity with carina; both dorsal and posterior faces flanked with high lamellae. In profile view, promesonotum high and dome-like; mesopleuron weakly differentiated into upper and lower portions; lower portion anteriorly and ventrally margined with carinae; metanotum separated from mesonotum and propodeum by deep furrows. Propodeum higher than long, with dorsal outline almost flat, armed at posterior angle with an angulate tooth posterodorsally directed; posterior margin lamellate; propodeal lobe triangular. Propodeal spiracle round and situated slightly above mid-height of sclerite, and slightly posterior to mid-length of sclerite; distance from propodeal spiracle to posterodorsal corner of propodeum (<abbrev xlink:title="propodeal spine length" id="ABBRID0E1CAG">PSL</abbrev> 0.13–0.14) about four times the diameter of spiracle (<abbrev xlink:title="propodeal spiracle diameter" id="ABBRID0E5CAG">PSD</abbrev> 0.03–0.04), and distance to declivity (<abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0ECDAG">DPSD</abbrev> 0.06–0.07) about twice the diameter of spiracle.</p>
            <p>Petiolar node, in dorsal view, wider than long (<abbrev xlink:title="petiolar node width" id="ABBRID0EIDAG">PTW</abbrev> 0.18–0.19, <abbrev xlink:title="petiolar node length" id="ABBRID0EMDAG">PNL</abbrev> 0.09–0.10); petiolar peduncle long, longer than wide, widened posteriad. Postpetiolar node nearly rectangular in outline, wider than long, slightly wider than petiolar node (<abbrev xlink:title="postpetiole width" id="ABBRID0EQDAG">PPW</abbrev> 0.19–0.21, <abbrev xlink:title="petiolar node width" id="ABBRID0EUDAG">PTW</abbrev> 0.18–0.19). Petiole, in profile view, longer than high (<abbrev xlink:title="petiole length" id="ABBRID0EYDAG">PTL</abbrev> 0.26–0.28, <abbrev xlink:title="petiolar node height" id="ABBRID0E3DAG">PTH</abbrev> 0.18–0.19) and with long peduncle; ventral outline of petiole nearly flat, anteriorly with subpetiolar process that is reduced to a small convexity; node differentiated from peduncle by rather distinct flection; node apically blunt, with posterior outline convex and with similar inclination as anterior outline. Postpetiolar node rounded and lower than petiolar node.</p>
            <p>Dorsal surface of head (frons and vertex) finely reticulate-rugose, with interspaces closely punctate; irregular short rugulae present just behind posterior margin of cephalic shield; posterior one-fourth of head dorsum with irregular transverse rugulae especially around horns; horn with longitudinal rugulae which extend to tip; dorsolateral face of head areolate-rugose. Upper two-thirds of phragmotic shield (mainly clypeus) finely shagreened and mat, with irregularly arranged stout rugulae. In profile, lateral face of head closely and minutely punctate; posterolateral portion of cephalic dorsum finely areolate-rugose. Dorsum of mandibles (constituting lower one-third of phragmotic shield) with similar surface sculpture as in upper portion of phragmotic shield. Pronotum almost entirely smooth and shiny, with polka-dot pattern seen through transparent cuticle; narrowed anterior portion (neck) and anterolateral area closely punctate and mat to coriaceous. Mesonotum superficially punctate and rather shiny; mesopleuron generally smooth and shiny. Metanotum (elevated narrowly transverse sclerite) sparsely punctate and shiny; furrows flanking it anteriorly and posteriorly minutely punctate and mat; metapleuron entirely closely puncto-reticulate. Propodeum entirely puncto-reticulate including spines and lateral lamellae on declivity. Petiole and postpetiole closely puncto-reticulate except for posterior face of postpetiole weakly rugose and dorsal central portion of postpetiole smooth and shiny. Gaster smooth and shiny, with anterior narrowed portion striate.</p>
            <p>Dorsal face of head with erect hairs sparsely, mixed with sparse appressed pubescence; anterior portion or cranium just behind phragmotic shield with three pairs of erect hairs (two pairs on supraclypeal area anteriorly and laterally, and one pair near antennal socket); lateral margin of head with few long hairs, and posterolateral corner of head around horn with suberect to curved hairs of various length. No visible hairs on cephalic shield. Outer margin of mandible with inconspicuous short appressed hairs; ventral face with erect or suberect hairs. Scape with appressed short hairs, without erect hairs. Promesonotum with less than ten long and fine erect hairs and sparse appressed pubescence. Propodeum often with a pair of suberect hairs. Petiole and postpetiole each with a pair of long suberect hairs and some decumbent hairs. Gaster with rather long appressed pubescence and long suberect hairs sparsely. Tibiae with decumbent hairs, without standing hairs. Body yellowish ferruginous; legs and parts of gaster, slightly lighter.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0ECEAG">
            <title>Description of minor worker.</title>
            <p>Head, in full-face view, long, about 1.2 times as long as wide (<abbrev xlink:title="cephalic index" id="ABBRID0EIEAG">CI</abbrev> 81–84), weakly subrectangular and slightly narrowed anteriorly; posterior margin almost straight, posterolateral corners rounded, and lateral margins convex. Frontal lobes elongate, widened anteriad, covering antennal sockets; frontal carinae short, as long as frontal lobe, diverging posteriad, with anterior two-fifths thickened. Frontal triangle distinct, shallowly depressed. Clypeus posteriorly extends to level of anterior extremity of frontal carina, between lobes almost as wide as lobe. Median disk of clypeus with fine lateral carinae that diverge anteriad; anterior margin of clypeus widely shallowly emarginate, with roundly angulate lateral corners that are lamellate; the lamella extends entire length of lateral margin of median disk of clypeus. Eyes present, consisting of one ommatidium (<abbrev xlink:title="eye length" id="ABBRID0EMEAG">EL</abbrev> 0.01). Mandibles with five teeth. Antennae with nine segments; scape fails to reach posterior margin of head (<abbrev xlink:title="head length" id="ABBRID0EQEAG">HL</abbrev> 0.36–0.38, <abbrev xlink:title="scape length" id="ABBRID0EUEAG">SL</abbrev> 0.23–0.24, <abbrev xlink:title="scape index" id="ABBRID0EYEAG">SI</abbrev> 61–65).</p>
            <p>In dorsal view, pronotum large, longer than wide; main sclerite round with anterior margin nearly straight and lateral margin roundly convex; prescutum (neck) of pronotum rather wide, slightly less than two-thirds as wide as main sclerite. Mesonotum not clearly demarcated from pronotum, narrower than the latter, tapered posteriad. Metanotal groove distinct, transverse, laterally margined with carinae. Propodeal dorsum longer than wide, widened posteriad, with long spines slightly directed outwardly. In profile view, promesonotum moderately and roundly convex dorsally. Pronotum with distinctly defined prescutum. Mesopleuron clearly differentiated from pronotum and metapleuron, divided into upper and lower portions by type of sculpture; lower portion anteriorly margined with distinct lamella. Metanotal groove impressed. Propodeum not clearly demarcated from metapleuron, about 1.5 times as high as long, with distinct dorsal face declining posteriorly, spines long and acute, posterodorsally directed and ventrally equipped with lamella that continues along posterior margin of declivity; the latter portion of lamella inflated below. Propodeal lobe triangular; spiracle rounded and situated slightly above mid-height of sclerite, and close to posterior margin of propodeum; distance from propodeal spiracle to posterodorsal corner of propodeum (<abbrev xlink:title="propodeal spine length" id="ABBRID0E5EAG">PSL</abbrev> 0.07–0.09) much longer than diameter of spiracle (<abbrev xlink:title="propodeal spine length" id="ABBRID0ECFAG">PSL</abbrev> 0.01–0.02), and distance to declivity (<abbrev xlink:title="distance from propodeal spiracle to declivity" id="ABBRID0EGFAG">DPSD</abbrev> 0.01–0.02) as large as diameter of spiracle.</p>
            <p>With waist in dorsal view, petiolar node as wide as long (<abbrev xlink:title="petiolar node width" id="ABBRID0EMFAG">PTW</abbrev> 0.06–0.07, <abbrev xlink:title="petiolar node length" id="ABBRID0EQFAG">PNL</abbrev> 0.06–0.07), and postpetiolar node slightly wider than petiolar node (<abbrev xlink:title="postpetiole width" id="ABBRID0EUFAG">PPW</abbrev> 0.08–0.09, <abbrev xlink:title="petiolar node width" id="ABBRID0EYFAG">PTW</abbrev> 0.06–0.07). Postpetiole wider than long, slightly wider than petiole. In profile view, petiole longer than high (<abbrev xlink:title="petiole length" id="ABBRID0E3FAG">PTL</abbrev> 0.15–0.16, <abbrev xlink:title="petiolar node height" id="ABBRID0EAGAG">PTH</abbrev> 0.10–0.11) with relatively long peduncle; sternite ventrally equipped with lamella over its length and has almost straight outline. Petiolar node demarcated from anterior peduncle by distinct bent; its dorsum with flat outline, and longer than anterior slope and as long as posterior slope; posterior margin of node convex. Subpetiolar process forming a blunt denticle, its height measuring almost same as diameter of propodeal spiracle. Postpetiole much smaller and lower than petiole and globular; node weakly convex dorsally. Anterolateral corners of first gastral tergite distinctly tuberculate.</p>
            <p>Dorsal surface of head including clypeus principally smooth, except for frontal lobe, lateral transverse area of clypeus and area between antennal socket and outer margin of head that are longitudinally rugose; anterior slope of clypeus punctate. Lateral and ventral faces of head principally smooth and shiny. Mandibles nearly smooth on dorsal face, superficially sculptured on laterobasal portion. Promesonotum smooth and shiny, except for prescutum (neck) of pronotum and posterior extremity of mesonotum puncto-reticulate. Lower portions of mesopleuron and metapleuron punctate; upper portions of mesopleuron and metapleuron smooth. Dorsal surface and posterior declivity (upper half) of propodeum densely punctate (punctures larger than those on mesopleuron); lower half of declivity smooth. Petiole and postpetiole densely punctate, except for dorsal central portion of postpetiole very weakly sculptured and shiny. Gastral tergites smooth and shiny except for presclerite of first tergite densely sculptured.</p>
            <p>Dorsum of head with sparse appressed/decumbent short hairs and a few erect hairs; lateral margin and posterior margin of head sparsely with erect/suberect hairs. Scape with short decumbent hairs. Outer margin of mandible with decumbent hairs. Mesosoma with four pairs of erect hairs, one pair on pronotum, two pairs on mesonotum, and one pair on propodeum. Petiole and postpetiole each with two pairs of erect/suberect hairs; gaster with sparse suberect hairs. Femora and tibiae with decumbent hairs. Color yellow.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="remarks" id="SECID0EGGAG">
            <title>Remarks.</title>
            <p>This species belongs to the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="acutispina">acutispina</tp:taxon-name-part></tp:taxon-name></italic> group. In the major worker it is easily distinguished from the other Asian members of the genus by the combination of the following conditions: the anteriorly truncated head, posterolateral corner of head with a horn, 9-segmented antennae, distinct metanotal disc, deep groove in front of the disc, developed propodeal spines, and laterally lamellate propodeal declivity. Minor workers characteristically have lamellae on the venter of the petiole and on the lateral margins of the propodeal declivity (the lamella on the propodeal declivity involves propodeal spine and extends anteriad up to the anterior margin of the propodeal dorsum as a low carina), and sparse erect hairs on the body.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="habitat" id="SECID0EXGAG">
            <title>Habitat.</title>
            <p>The type series was collected from leaf litter in a lowland dry evergreen forest (Fig. <xref ref-type="fig" rid="F4">4</xref>).</p>
            <fig id="F4" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/jhr.91.82490.figure4</object-id>
              <object-id content-type="arpha">7791A8A9-06BF-5888-BA84-73BFA5456DB4</object-id>
              <label>Figure 4.</label>
              <caption>
                <p>Forest habitats on lowland dry evergreen forest, Siem Reap Province, Cambodia, the type locality of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (S. Hosoishi).</p>
              </caption>
              <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707303.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/707303</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0EZHAG">
            <title>Distribution.</title>
            <p>This species is only known from the type locality in Cambodia.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0E5HAG">
            <title>Etymology.</title>
            <p>The species name refers to the anteriorly truncated head of the major worker subcaste.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="SECID0EDIAG">
      <title>Discussion</title>
      <p>This is the first confirmed record of a truncated phragmotic head in the major worker of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic>. Among Asian <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> fauna, the three species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic> (Forel, 1913), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> (Forel, 1902) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name></italic> (Sheela &amp; Narendran, 1997) are known as the phragmotic species with a depressed cephalic shield. <xref ref-type="bibr" rid="B13">Forel (1913)</xref> suggested that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic> is similar or conspecific to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic>, but concluded that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> is different in having the developed rugulae on head, much less concave frons and occipital region not emarginated. In his paper, <xref ref-type="bibr" rid="B13">Forel (1913)</xref> established the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="infraspecific-rank">subgenus</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subgenus" reg="Lecanomyrma">Lecanomyrma</tp:taxon-name-part></tp:taxon-name> of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidologeton">Pheidologeton</tp:taxon-name-part></tp:taxon-name></italic> based on a depressed cephalic shield in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic> (as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidologeton">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic>) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> (as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidologeton">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic>). <xref ref-type="bibr" rid="B7">Emery (1924)</xref> transferred the two species to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Aneleus">Aneleus</tp:taxon-name-part> (<tp:taxon-name-part taxon-name-part-type="subgenus" reg="Lecanomyrma">Lecanomyrma</tp:taxon-name-part>)</tp:taxon-name>. <xref ref-type="bibr" rid="B8">Ettershank (1966)</xref> treated them as members of the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Oligomyrmex">Oligomyrmex</tp:taxon-name-part></tp:taxon-name></italic>, then <xref ref-type="bibr" rid="B9">Fernández (2004)</xref> synonymized the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Oligomyrmex">Oligomyrmex</tp:taxon-name-part></tp:taxon-name></italic> under the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic>.</p>
      <p>The Afrotropical species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> has a dimorphic major subcaste, one with the phragmotic head and the other with the regular head shape (<xref ref-type="bibr" rid="B11">Fischer et al. 2015</xref>). The regular major workers are so far unknown in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic>. Although our material is limited, all the major workers examined had an anteriorly truncated head. The regular major workers are also unknown for South Asian species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name></italic>. Those incomplete collections are possibly due to the rarity of the phragmotic species (<xref ref-type="bibr" rid="B11">Fischer et al. 2015</xref>). The nest series samples might reveal the occurrence of the regular major workers in Asian phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species in the future.</p>
      <p>The phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species seem to share several morphological trends (Table <xref ref-type="table" rid="T1">1</xref>), but separated based on the antennal counts into three geographical members: Southeast Asian (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic>), South Asian (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic>) and Afrotropical (the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> species clade: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="elmenteitae">elmenteitae</tp:taxon-name-part></tp:taxon-name></italic> (Patrizi, 1948); <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lilith">lilith</tp:taxon-name-part></tp:taxon-name></italic> Fischer, Azorsa &amp; Hita Garcia, 2015; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> Fischer, Azorsa &amp; Hita Garcia, 2015). The number of antennal segments are considered one of the phylogenetically important characters in ants (e.g., <xref ref-type="bibr" rid="B22">Sparks et al. 2019</xref>). This suggests that the phragmotic mode of life evolved independently in different lineages, although a comprehensive phylogeny of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> has not been presented.</p>
      <table-wrap id="T1" position="float" orientation="portrait">
        <label>Table 1.</label>
        <caption>
          <p>Character states of the phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> major workers. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> is not shown here because the major worker caste is unknown.</p>
        </caption>
        <table id="TID0EFQAG" rules="all">
          <tbody>
            <tr>
              <th rowspan="1" colspan="1">Species</th>
              <th rowspan="1" colspan="1">Antennal segments</th>
              <th rowspan="1" colspan="1">Horn on posterolateral corner of head</th>
              <th rowspan="1" colspan="1">Metanotum</th>
              <th rowspan="1" colspan="1">Propodeal spines</th>
              <th rowspan="1" colspan="1">Propodeal declivity</th>
              <th rowspan="1" colspan="1">Reference</th>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">9</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">lamellate</td>
              <td rowspan="1" colspan="1">This study</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">11</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">?</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">?</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B21">Sheela and Narendran (1997)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">11</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">?present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">?lamellate</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B13">Forel (1913)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">10</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">lamellate</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="elmenteitae">elmenteitae</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">10</td>
              <td rowspan="1" colspan="1">absent</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">lamellate</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lilith">lilith</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">10</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">present</td>
              <td rowspan="1" colspan="1">lamellate</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
      <p>The truncated head shape in the major worker of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. represents a type of phragmosis seen in ants, but is different from those found in the Afrotropical (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="elmenteitae">elmenteitae</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lilith">lilith</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic>) and South Asian <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name></italic>). The truncated area is composed of clypeus and mandibles in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic>, while it is the depressed dorsal surface of the cranium, clypeus and mandibles in the above-mentioned species. As mentioned earlier, phragmosis has been reported from several ant genera (Figs <xref ref-type="fig" rid="F5">5</xref>, <xref ref-type="fig" rid="F6">6</xref>, Table <xref ref-type="table" rid="T2">2</xref>). The phragmotic head in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">C.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> is similar to those in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Crematogaster">Crematogaster</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cylindriceps">cylindriceps</tp:taxon-name-part></tp:taxon-name></italic> W. M. Wheeler, 1927, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamia">lamia</tp:taxon-name-part></tp:taxon-name></italic> W. M. Wheeler, 1901, and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tetraponera">Tetraponera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> Ward, 2006 rather than those in other phragmotic <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part></tp:taxon-name></italic> species.</p>
      <fig id="F5" position="float" orientation="portrait">
        <object-id content-type="doi">10.3897/jhr.91.82490.figure5</object-id>
        <object-id content-type="arpha">847E388C-4862-524B-BAA2-C6C246BD09D9</object-id>
        <label>Figure 5.</label>
        <caption>
          <p>Phragmotic ants <bold>A</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name></italic> worker (Lubertazzi) <bold>B</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name></italic> queen, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0908890">CASENT0908890</ext-link> (Zach Lieberman) <bold>C</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name></italic> worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0908888">CASENT0908888</ext-link> (Zach Lieberman) <bold>D</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> phragmotic major worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0709550">CASENT0709550</ext-link>. Image from <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref><bold>E</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="elmenteitae">elmenteitae</tp:taxon-name-part></tp:taxon-name></italic> phragmotic major worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0217087">CASENT0217087</ext-link> (Erin Prado) <bold>F</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lilith">lilith</tp:taxon-name-part></tp:taxon-name></italic> phragmotic major worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0709545">CASENT0709545</ext-link>. Image from <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref><bold>G</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Crematogaster">Crematogaster</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cylindriceps">cylindriceps</tp:taxon-name-part></tp:taxon-name></italic> queen. Image from <xref ref-type="bibr" rid="B32">Yamane et al. (2011)</xref><bold>H</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name></italic> worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0619464">CASENT0619464</ext-link> (Jeremy Pilllow) <bold>I</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamia">lamia</tp:taxon-name-part></tp:taxon-name></italic> worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0104664">CASENT0104664</ext-link> (April Nobile). <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://www.AntWeb.org">www.AntWeb.org</ext-link>.</p>
        </caption>
        <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707304.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/707304</uri>
        </graphic>
      </fig>
      <fig id="F6" position="float" orientation="portrait">
        <object-id content-type="doi">10.3897/jhr.91.82490.figure6</object-id>
        <object-id content-type="arpha">39EC5A49-1D66-57A1-B50A-9C8886ECB5BC</object-id>
        <label>Figure 6.</label>
        <caption>
          <p>Phragmotic ants (continued) <bold>A</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="embolopyx">embolopyx</tp:taxon-name-part></tp:taxon-name></italic> queen. Arrow indicates the phragmotic shield on gaster. Modified from Brown (1968) <bold>B</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blepharidatta">Blepharidatta</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="conops">conops</tp:taxon-name-part></tp:taxon-name></italic> worker. Image from Brandão et al. (2015) <bold>C</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobostruma">Colobostruma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leae">leae</tp:taxon-name-part></tp:taxon-name></italic> holotype queen (Steve Shattcuk) <bold>D</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Cephalotes">Cephalotes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="atratus">atratus</tp:taxon-name-part></tp:taxon-name></italic> soldier worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0178627">CASENT0178627</ext-link> (April Nobile) <bold>E</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobopsis">Colobopsis</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cerberula">cerberula</tp:taxon-name-part></tp:taxon-name></italic> major worker ,<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0104765">CASENT0104765</ext-link> (April Nobile) <bold>F</bold><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tetraponera">Tetraponera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name></italic> major worker, <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://data.antweb.org/specimen/CASENT0136420">CASENT0136420</ext-link> (April Nobile). <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://www.AntWeb.org">www.AntWeb.org</ext-link>.</p>
        </caption>
        <graphic xlink:href="journal_of_hymenoptera_research-91-357_article-82490__-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_707305.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/707305</uri>
        </graphic>
      </fig>
      <table-wrap id="T2" position="float" orientation="portrait">
        <label>Table 2.</label>
        <caption>
          <p>Body parts used for phragmosis in ants.</p>
        </caption>
        <table id="TID0ES2AG" rules="all">
          <tbody>
            <tr>
              <th rowspan="1" colspan="1">Subfamily</th>
              <th rowspan="1" colspan="1">Taxon</th>
              <th rowspan="1" colspan="1">Caste</th>
              <th rowspan="1" colspan="1">Body parts used for phragmosis</th>
              <th rowspan="1" colspan="1">Distribution</th>
              <th rowspan="1" colspan="1">Reference</th>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Clypeus, mandible</td>
              <td rowspan="1" colspan="1">Southeast Asia (Cambodia)</td>
              <td rowspan="1" colspan="1">This study</td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="nayana">nayana</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">South Asia (India)</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B21">Sheela and Narendran (1997)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamellifrons">lamellifrons</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">queen</td>
              <td rowspan="1" colspan="1">Cranium, clypeus</td>
              <td rowspan="1" colspan="1">South Asia (India)</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B12">Forel (1902</xref>, <xref ref-type="bibr" rid="B13">1913</xref>), <xref ref-type="bibr" rid="B7">Emery (1924)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="butteli">butteli</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">South Asia (Sri Lanka)</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B13">Forel (1913)</xref>, <xref ref-type="bibr" rid="B7">Emery (1924)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">Africa (Kenya)</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="elmenteitae">elmenteitae</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">Africa (Kenya)</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Carebara">Carebara</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lilith">lilith</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">Africa (Ivory coast)</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B11">Fischer et al. (2015)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Crematogaster">Crematogaster</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="cylindriceps">cylindriceps</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">queen</td>
              <td rowspan="1" colspan="1">Clypeus, mandible</td>
              <td rowspan="1" colspan="1">Southeast Asia (Malaysia)</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B29">Wheeler (1927)</xref>, <xref ref-type="bibr" rid="B32">Yamane et al. (2011)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="colobopsis">colobopsis</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker, queen</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">New World tropics</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B18">Mann (1916)</xref>, <xref ref-type="bibr" rid="B30">Wilson (2003)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lamia">lamia</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker</td>
              <td rowspan="1" colspan="1">Clypeus, mandible</td>
              <td rowspan="1" colspan="1">United States, Mexico</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B28">Wheeler (1901)</xref>, <xref ref-type="bibr" rid="B30">Wilson (2003)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pheidole">Pheidole</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="embolopyx">embolopyx</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">queen</td>
              <td rowspan="1" colspan="1">Gaster</td>
              <td rowspan="1" colspan="1">South America</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B6">Brown Jr (1968)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Blepharidatta">Blepharidatta</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="conops">conops</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">queen</td>
              <td rowspan="1" colspan="1">Cranium, pronotum</td>
              <td rowspan="1" colspan="1">New World tropics</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B5">Brandão et al. (2001)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Colobostruma">Colobostruma</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="leae">leae</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">worker, queen</td>
              <td rowspan="1" colspan="1">Cranium, clypeus, mandible</td>
              <td rowspan="1" colspan="1">Australia</td>
              <td rowspan="1" colspan="1"><xref ref-type="bibr" rid="B29">Wheeler (1927)</xref>, <xref ref-type="bibr" rid="B20">Shattuck (2000)</xref></td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Myrmicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="genus" reg="Cephalotes">Cephalotes</tp:taxon-name-part>
                  </tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker, queen</td>
              <td rowspan="1" colspan="1">Cranium</td>
              <td rowspan="1" colspan="1">New World tropics</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B19">Oliveira et al. (2021)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Formicinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name>
                    <tp:taxon-name-part taxon-name-part-type="genus" reg="Colobopsis">Colobopsis</tp:taxon-name-part>
                  </tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker, queen</td>
              <td rowspan="1" colspan="1">Cranium (malar), clypeus, mandible</td>
              <td rowspan="1" colspan="1">Palearctic, Oriental, Australian, New World (part)</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B26">Ward et al. (2016)</xref>
              </td>
            </tr>
            <tr>
              <td rowspan="1" colspan="1">
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="subfamily">Pseudomyrmecinae</tp:taxon-name-part>
                </tp:taxon-name>
              </td>
              <td rowspan="1" colspan="1">
                <italic>
                  <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tetraponera">Tetraponera</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="phragmotica">phragmotica</tp:taxon-name-part></tp:taxon-name>
                </italic>
              </td>
              <td rowspan="1" colspan="1">major worker, queen</td>
              <td rowspan="1" colspan="1">Clypeus, mandible</td>
              <td rowspan="1" colspan="1">Madagascar</td>
              <td rowspan="1" colspan="1">
                <xref ref-type="bibr" rid="B25">Ward (2006)</xref>
              </td>
            </tr>
          </tbody>
        </table>
      </table-wrap>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>We would like to thank Phourin Chhang and Choeung Hong Narith (Forestry Administration of Cambodia, Phnom Penh, Cambodia) for providing permission to conduct research in Cambodia, issuing a specimen export permit to Kyushu University, Japan, and suggesting the type depository of new species. We also express our thanks to Tsuyoshi Kajisa (Kagoshima University), Nobuya Mizoue (Kyushu University), Tetsukazu Yahara (Kyushu University) for helping our field surveys in this study. We would like to thank Frank Azorsa, Georg Fischer and Rodolfo Probst for valuable comments on the manuscript. We would like to thank Editage (<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://www.editage.com">www.editage.com</ext-link>) for English language editing. We would like to thank ANeT members for encouragement. This work was supported in part by JSPS KAKENHI (Grant-in-Aid for Scientific Research (C) Grant Number 21K05616, 19K06824, and Global COE Program (Center of Excellence for Asian Conservation Ecology as a basis of Human-Nature Mutualism), MEXT, Japan.</p>
    </ack>
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