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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">65</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:EC092FB7-29CB-5510-A337-7A4057D255E5</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:75EDC164-A8BB-4D1B-88FC-F9FE7054E1A3</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Evolutionary Systematics</journal-title>
        <abbrev-journal-title xml:lang="en">EvolSyst</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">2535-0730</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/evolsyst.8.124285</article-id>
      <article-id pub-id-type="publisher-id">124285</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Tetrigidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Biogeography</subject>
          <subject>Nomenclature</subject>
          <subject>Taxonomy</subject>
        </subj-group>
        <subj-group subj-group-type="geological_era">
          <subject>Cenozoic</subject>
        </subj-group>
        <subj-group subj-group-type="geographical_area">
          <subject>Ecuador</subject>
          <subject>French Guyana/Guyane</subject>
          <subject>Peru</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>﻿Description of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> from Ecuador with notes on the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Orthoptera</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name>)</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Kasalo</surname>
            <given-names>Niko</given-names>
          </name>
          <email xlink:type="simple">niko.kasalo5@gmail.com</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-3139-6349</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Husemann</surname>
            <given-names>Martin</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-5536-6681</uri>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>van de Kamp</surname>
            <given-names>Thomas</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0001-7390-1318</uri>
          <xref ref-type="aff" rid="A3">3</xref>
          <xref ref-type="aff" rid="A4">4</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Skejo</surname>
            <given-names>Josip</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-2554-4499</uri>
          <xref ref-type="aff" rid="A5">5</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Laboratory of Evolutionary Genetics, Ruđer Bošković Institute, Bijenička cesta 54, HR-10000 Zagreb, Croatia</addr-line>
        <institution>Laboratory of Evolutionary Genetics, Ruđer Bošković Institute</institution>
        <addr-line content-type="city">Zagreb</addr-line>
        <country>Croatia</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Staatliches Museum für Naturkunde Karlsruhe, Erbprinzenstraße 13, D-76133 Karlsruhe, Germany</addr-line>
        <institution>Staatliches Museum für Naturkunde Karlsruhe</institution>
        <addr-line content-type="city">Karlsruhe</addr-line>
        <country>Germany</country>
      </aff>
      <aff id="A3">
        <label>3</label>
        <addr-line content-type="verbatim">Karlsruhe Institute of Technology (KIT), Institute for Photon Science and Synchrotron Radiation (IPS), Hermann-von-Helmholtz-Platz 1, D-76344 Eggenstein-Leopoldshafen, Karlsruhe, Germany</addr-line>
        <institution>Karlsruhe Institute of Technology (KIT), Institute for Photon Science and Synchrotron Radiation (IPS)</institution>
        <addr-line content-type="city">Karlsruhe</addr-line>
        <country>Germany</country>
      </aff>
      <aff id="A4">
        <label>4</label>
        <addr-line content-type="verbatim">Karlsruhe Institute of Technology (KIT), Laboratory for Applications of Synchrotron Radiation, Kaiserstr. 12, D-76131 Karlsruhe, Germany</addr-line>
        <institution>Karlsruhe Institute of Technology (KIT), Laboratory for Applications of Synchrotron Radiation</institution>
        <addr-line content-type="city">Karlsruhe</addr-line>
        <country>Germany</country>
      </aff>
      <aff id="A5">
        <label>5</label>
        <addr-line content-type="verbatim">University of Zagreb, Faculty of Science, Department of Biology, Evolution Lab, Horvatovac 102a, HR-10000 Zagreb, Croatia</addr-line>
        <institution>University of Zagreb</institution>
        <addr-line content-type="city">Zagreb</addr-line>
        <country>Croatia</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Niko Kasalo (<email xlink:type="simple">nkasalo@irb.hr</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Academic editor: Oliver Hawlitschek</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2024</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>28</day>
        <month>05</month>
        <year>2024</year>
      </pub-date>
      <volume>8</volume>
      <issue>1</issue>
      <fpage>119</fpage>
      <lpage>125</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/9B98BC33-A31F-5714-8578-DCCF4EADFDF6">9B98BC33-A31F-5714-8578-DCCF4EADFDF6</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/7583B1DD-2B0C-49ED-A68C-7C02C1F1AECC">7583B1DD-2B0C-49ED-A68C-7C02C1F1AECC</uri>
      <history>
        <date date-type="received">
          <day>30</day>
          <month>03</month>
          <year>2024</year>
        </date>
        <date date-type="accepted">
          <day>07</day>
          <month>05</month>
          <year>2024</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Niko Kasalo, Martin Husemann, Thomas van de Kamp, Josip Skejo</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/7583B1DD-2B0C-49ED-A68C-7C02C1F1AECC</self-uri>
      <abstract>
        <label>﻿Abstract</label>
        <p><tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part>
          </tp:taxon-name> Hancock, 1909, endemic to the Neotropics, is one of the most ill-known subfamilies of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name>. Until now, there have not been any hypothesis on its relationship with other subfamilies. <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> comprise only two genera, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic> Hancock, 1909 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> Bolívar, 1906, both with an arduous taxonomic history. Here, we describe a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> from Ecuador, and compare it with its congeners: <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> from Peru and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> from French Guiana. We found the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> to most closely resemble the Neotropical subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Metrodorinae</tp:taxon-name-part></tp:taxon-name>, meaning that the two are likely related. We also provide a 3D scan of the holotype, a first for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name>. The 3D model contains all the important taxonomic characters and much more data could be extracted in the future using artificial intelligence-assisted approaches.</p>
      </abstract>
      <kwd-group>
        <label>Key Words</label>
        <kwd>3D scan</kwd>
        <kwd>Neotropics</kwd>
        <kwd>taxonomy</kwd>
        <kwd>pygmy hoppers</kwd>
        <kwd>grouse hopper</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="﻿Introduction" id="SECID0E4H">
      <title>﻿Introduction</title>
      <p><tp:taxon-name>
          <tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part>
        </tp:taxon-name> are a relatively specious orthopteran family with around 2000 described species (<xref ref-type="bibr" rid="B6">Cigliano et al. 2024</xref>). The family is currently undergoing revision at all taxonomic levels, with papers being regularly published on taxa in all biogeographical regions (<xref ref-type="bibr" rid="B6">Cigliano et al. 2024</xref>). The Neotropical region is especially understudied–only a few recent studies have introduced some brief taxonomic acts (<xref ref-type="bibr" rid="B14">Itrac-Bruneau and Doucet 2022</xref>, <xref ref-type="bibr" rid="B15">2023</xref>; <xref ref-type="bibr" rid="B16">Kasalo et al. 2022</xref>, <xref ref-type="bibr" rid="B17">2023</xref>).</p>
      <p>Among the many problematic taxa, the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> Hancock, 1909 is perhaps the most elusive one. It comprises only two genera, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic> Hancock, 1909 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> Bolívar, 1906, the specimens of which are extremely rarely observed (<xref ref-type="bibr" rid="B16">Kasalo et al. 2022</xref>; <xref ref-type="bibr" rid="B15">Itrac-Bruneau and Doucet 2023</xref>). Furthermore, it is the only tetrigid subfamily endemic to the Neotropical biogeographic region. Likely, also due to its rarity, there is a complete absence of information on its position in the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name> phylogenetic tree.</p>
      <p>The genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> was briefly synonymized under <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chiriquia">Chiriquia</tp:taxon-name-part></tp:taxon-name></italic> Morse, 1900 (<xref ref-type="bibr" rid="B4">Cadena-Castañeda and Cardona 2015</xref>), but was quickly resurrected alongside a description of a new species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B5">Cadena-Castañeda et al. 2021</xref>). The genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic> has already been revised to the extent that the currently available data permits (Silva et al. 2019; <xref ref-type="bibr" rid="B16">Kasalo et al. 2022</xref>), leaving only <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> to be reevaluated in light of recent taxonomic advances.</p>
      <p>One important step to facilitate future taxonomic revisions and make them more independent of the geographic position of the researchers is enhancing digitalization efforts. The digitalization of type specimens is of special importance since it allows quick and easy access to the material without risking damaging specimens and without consuming the curators’ time. However, the vast majority of insect type specimens has not been photographed, and many of those that have been digitalized in this way only have dorsal views available, which severely limits their usefulness (<xref ref-type="bibr" rid="B20">Mertens et al. 2017</xref>; <xref ref-type="bibr" rid="B24">Ströbel et al. 2018</xref>). For proper taxonomic utilization, multiple views should be provided, and ideally, a scaled 3D model would be produced since it allows the specimen to be seen as it exists in reality, and more precise measurements can be taken, even without the actual specimen at hand (<xref ref-type="bibr" rid="B24">Ströbel et al. 2018</xref>). Unfortunately, even the simple photographic equipment is prohibitively expensive for many institutions that house type specimens, making well-photographed types rare and 3D scans nearly non-existent (<xref ref-type="bibr" rid="B20">Mertens et al. 2017</xref>; <xref ref-type="bibr" rid="B24">Ströbel et al. 2018</xref>). Until now, no 3D scans of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name> existed.</p>
      <p>Here, we describe <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov., a new species of this understudied genus and provide a 3D scan of the holotype, a first among <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name>. Furthermore, we review the status of the genus and provide the first hypothesis about the phylogenetic placement of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name>.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EMFAC">
      <title>﻿Materials and methods</title>
      <sec sec-type="﻿Basic photography and measurements" id="SECID0EQFAC">
        <title>﻿Basic photography and measurements</title>
        <p>The specimen was examined and photographed by Mathias Vielsäcker at SMNK using a Canon EOS R5 with either a Canon EF 100MM F/2.8L MACRO IS USM for the dorsal, ventral and lateral images, or a Laowa 25 mm f/2,8 Ultra Macro 2,5–5X for the images of smaller details. Stacked imaging was performed using a WeMacro rail. Images were then stacked using Helicon Focus.</p>
        <p>Measurements were made using ImageJ 1.53t software (<xref ref-type="bibr" rid="B22">Rueden et al. 2017</xref>). Taxonomy follows the OSF (<xref ref-type="bibr" rid="B6">Cigliano et al. 2024</xref>). Morphological terminology and measurements follow <xref ref-type="bibr" rid="B25">Tumbrinck (2014)</xref>, with some additional measurements. The following measurements were taken: body length (<abbrev xlink:title="body length" id="ABBRID0EDGAC">BL</abbrev>), pronotum length (<abbrev xlink:title="pronotum length" id="ABBRID0EHGAC">PL</abbrev>), pronotum width (shoulders) (<abbrev xlink:title="pronotum width" id="ABBRID0ELGAC">PWS</abbrev>), pronotum width (maximal) (<abbrev xlink:title="pronotum width" id="ABBRID0EPGAC">PWM</abbrev>), pronotum height (maximal) (<abbrev xlink:title="pronotum height" id="ABBRID0ETGAC">PHM</abbrev>), vertex width (<abbrev xlink:title="vertex width" id="ABBRID0EXGAC">VW</abbrev>), compound eye width (<abbrev xlink:title="compound eye width" id="ABBRID0E2GAC">CEW</abbrev>), antennal groove width (<abbrev xlink:title="antennal groove width" id="ABBRID0E6GAC">AGW</abbrev>), scutellum width (<abbrev xlink:title="scutellum width" id="ABBRID0EDHAC">SW</abbrev>), fore femur length (<abbrev xlink:title="fore femur length" id="ABBRID0EHHAC">FFL</abbrev>), fore femur width (<abbrev xlink:title="fore femur width" id="ABBRID0ELHAC">FFW</abbrev>), mid femur length (<abbrev xlink:title="mid femur length" id="ABBRID0EPHAC">MFL</abbrev>), mid femur width (<abbrev xlink:title="mid femur width" id="ABBRID0ETHAC">MFW</abbrev>), hind femur length (<abbrev xlink:title="hind femur length" id="ABBRID0EXHAC">HFL</abbrev>), hind femur width (<abbrev xlink:title="hind femur width" id="ABBRID0E2HAC">HFW</abbrev>), tegmen length (<abbrev xlink:title="tegmen length" id="ABBRID0E6HAC">TL</abbrev>), tegmen width (<abbrev xlink:title="tegmen width" id="ABBRID0EDIAC">TW</abbrev>). Nomenclature is in accordance with the rules of the International Code of Zoological Nomenclature (<xref ref-type="bibr" rid="B13">ICZN 1999</xref>).</p>
      </sec>
      <sec sec-type="﻿Synchrotron X-ray microtomography" id="SECID0ELIAC">
        <title>﻿Synchrotron X-ray microtomography</title>
        <p>The specimen was scanned in six height steps using a filtered polychromatic beam at the high-throughput tomography station at the Imaging Cluster of KIT Light Source. We employed an air-bearing rotary stage (RT150S, LAB Motion Systems) for sample rotation and a fast indirect detector system consisting of a 200 µm LuAG scintillator, a double objective white beam microscope (Optique Peter; <xref ref-type="bibr" rid="B7">Douissard et al. 2012</xref>) and a 12-bit pco.dimax high-speed camera (Excelitas PCO GmbH) with 2016 × 2016 pixels of 11 µm physical size. We employed a magnification of 2×, resulting in an effective pixel size of 6.11 µm. For each scan, we took 200 dark field images, 100 flat field images, and 3000 equiangularly spaced radiographic projections in a range of 180° with an exposure time of 30 ms. The control system concert (<xref ref-type="bibr" rid="B27">Vogelgesang et al. 2016</xref>) served for automated data acquisition and online reconstruction of tomographic slices for data quality assurance. The 3D tomographic reconstruction was performed by tofu (Faragó et al. 2017) and additionally included ring removal and 8-bit conversion.</p>
      </sec>
      <sec sec-type="﻿Post-processing of tomographic data" id="SECID0EZIAC">
        <title>﻿Post-processing of tomographic data</title>
        <p>The individual tomographic volumes were registered and merged in Amira 2022.2, which was also employed for pre-segmentation. Pre-segmented labels served as input for semi-automatic segmentation with Biomedisa (<xref ref-type="bibr" rid="B19">Lösel et al. 2020</xref>). The Biomedisa result was imported back into Amira and minor errors were corrected. The needle was digitally removed and the holes filled. The final label field was converted into a polygon mesh and exported as OBJ file. CINEMA 4D R20 was employed for smoothing the surface and for polygon reduction. The polygon-reduced model was imported into MeshLab, converted into U3D format and integrated into an interactive 3D PDF document (Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>).</p>
      </sec>
    </sec>
    <sec sec-type="﻿Results" id="SECID0EHJAC">
      <title>﻿Results</title>
      <sec sec-type="﻿Taxonomy" id="SECID0ELJAC">
        <title>﻿Taxonomy</title>
        <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">Orthoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Tetrigidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿Subfamily</label>
            <tp:taxon-name><object-id content-type="arpha">B74027FA-50EA-5A92-8068-FBECD2330FB1</object-id>
              <tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>Hancock, 1909</tp:taxon-authority>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0EOKAC">
            <title>Diagnosis.</title>
            <p>Frontal costa bifurcation in upper third of eye height. Antennal groove at bottom margin of eyes. Antennomeres flattened, apical one white-tipped. Vertex wider than an eye. First segment of anterior and middle tarsi a little elongated. Hind tarsus very long, first and third segment approximately equally long. Elongated slim ovipositor.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type genus" id="SECID0ETKAC">
            <title>Type genus.</title>
            <p><italic><tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part>
                </tp:taxon-name></italic> Hancock, 1909, type species <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="brevicristatus">brevicristatus</tp:taxon-name-part></tp:taxon-name></italic> Hancock, 1909.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0EPLAC">
            <title>Composition and distribution.</title>
            <p>Two genera, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic>, endemic to Southern and Central America.</p>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <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-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿Genus</label>
            <tp:taxon-name><object-id content-type="arpha">04EFBEB1-C937-595E-97B8-4FE3798886F8</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part>
            </tp:taxon-name>
            <tp:taxon-authority>Bolívar, 1906</tp:taxon-authority>
            <tp:nomenclature-citation-list>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Nephele">Nephele</tp:taxon-name-part>
                </tp:taxon-name>
                <comment>Bolívar, 1887: 252. (junior homonym of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Nephele">Nephele</tp:taxon-name-part></tp:taxon-name> Hübner, 1819);</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part>
                </tp:taxon-name>
                <comment>Bolívar, 1906: 392. (replacement name); <xref ref-type="bibr" rid="B18">Kirby 1910</xref>: 23. (catalogued); Günther, 1938: 307. (diagnosis); <xref ref-type="bibr" rid="B28">Yin et al. 1996</xref>: 899. (catalogued); <xref ref-type="bibr" rid="B4">Cadena-Castañeda and Cardona 2015</xref>: 477. (noted as a synonym of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chiriquia">Chiriquia</tp:taxon-name-part></tp:taxon-name>); Silva et al. 2019 (mentioned); <xref ref-type="bibr" rid="B5">Cadena-Castañeda et al. 2021</xref>: 357. (revision);</comment>
              </tp:nomenclature-citation>
              <tp:nomenclature-citation>
                <tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Gladiotettix">Gladiotettix</tp:taxon-name-part>
                </tp:taxon-name>
                <comment>Hancock, 1907: 40. (diagnosis in English, obsolete replacement name); <xref ref-type="bibr" rid="B3">Bruner 1910</xref>: 101. (Hancock’s descriptions of the species which was later assigned to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name>).</comment>
              </tp:nomenclature-citation>
            </tp:nomenclature-citation-list>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="Differential diagnosis" id="SECID0EIPAC">
            <title>Differential diagnosis.</title>
            <p>Antennal segments flattened, but oval (apical segments triangular in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic>). Low pronotal crest (medium to tall in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic>). Anterior and middle femora smooth (moderately tuberculated in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lophotettix">Lophotettix</tp:taxon-name-part></tp:taxon-name></italic>).</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type species" id="SECID0EEAAE">
            <title>Type species.</title>
            <p><italic><tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> (Bolívar, 1887)</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0EYAAE">
            <title>Composition and distribution.</title>
            <p><italic><tp:taxon-name>
                  <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> Cadena-Castañeda &amp; Tumbrinck, 2021, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> (Bolívar, 1887), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. The genus seems to be restricted to the northern half of South America.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="notes" id="SECID0ECCAE">
            <title>Note.</title>
            <p>We identified two female specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> from Peru at the Zoologische Staatssammlung München (ZSM), which we present in Fig. <xref ref-type="fig" rid="F1">1</xref> to aid in comparison with the new species. Additionally, we identified two male specimens of this species at the ZSM with the following label data: Peru, Prov. Huanuco / b. Yuyapichis, Stat. Panguana / <named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-74.600000,-9.616667]}" id="NCID0E1CAE">9°37'S, 74°36'W</named-content></named-content> Lux Wald / Baum liegend, 20.9.–7.10.2013 / leg. Burmeister.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="arpha">7C93E61B-41FD-54DD-B03A-8C88D5027B6A</object-id>
              <label>Figure 1.</label>
              <caption>
                <p><italic><tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> specimens deposited in ZSM. <bold>A.</bold> The locality where the specimens were collected; <bold>B.</bold> Label, same for both specimens; <bold>C.</bold> Two female specimens depicted in lateral, frontal, and dorsal views. All photos are to the same scale except the frontal views. Scale bar: 1mm.</p>
              </caption>
              <graphic xlink:href="evolsyst-08-119-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1056784.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1056784</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
        </tp:taxon-treatment>
        <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-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <label>﻿</label>
            <tp:taxon-name><object-id content-type="arpha">A73734B3-4D61-5409-847B-1658736AA67E</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">https://zoobank.org/6047D7A1-7F3F-40FF-BDD2-4E2B67592A16</object-id>
            </tp:taxon-name>
            <tp:taxon-authority>Kasalo, Husemann &amp; Skejo</tp:taxon-authority>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F2">Fig. 2</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0ENEAE">
            <title>Diagnosis.</title>
            <p>Easily separated from the congeners by the following set of characters: (i) median carina forming a low elongated crest which extends from between the prozonal carinae to the base of hind legs; (ii) vertex a little less than two eyes wide; (iii) brown body without distinct patterns; (iv) rounded pulvilli of hind tarsi; (v) female subgenital plate oval with moderately protruding triangular apex; (vi) alae dark brown with venation of the same color. Differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> by characters iii, iv, v, and vi.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0E5EAE">
            <title>Description.</title>
            <p>Macropronotal. In anterior view, top of vertex approximately at level of upper margin of eyes. Vertex a little less than two eyes wide. Frontal costa bifurcation in upper third of eye height. Facial carinae forming long oval shape. Paired ocelli approximately at half of eye height. Midline of antennal groove at level of bottom margin of eyes. Antennae composed of 13 visible segments; apical six flattened, last segment white. In dorsal view, vertex of isosceles trapezoidal shape. Medial carina visible in anterior half of vertex. Lateral carinae in form of small tubercles. Prozonal carinae as long as eye as seen in dorsal view, strongly convergent caudad. Median carina visible throughout length of pronotum, forming low but long crest in lateral view (extending from between prozonal carinae to base of hind legs). Pronotal surface covered in small tubercles and carinulae. Humeral angles wide and blunt. Lateral lobes rectangular, projected laterally, with thin white border. Tegmina large and oval. Alae reaching pronotal apex; dark brown with dark brown venation. Anterior and middle femora long and smooth. Anterior and middle femora long and smooth. Anterior and middle tarsi with long first segment and extremely long second segment. Hind femur robust, with two tubercles on dorsal margin; moderately sized sharp antegenicular tooth, genicular tooth barely visible. Hind tibia thin, serrated along caudal margin. Hind tarsus long; first segment extremely long, with three elongated pulvilli; third segment missing in the holotype. Ovipositor extremely elongated, serrated. Base of subgenital plate with triangular notch; shallow stitch line extending from it throughout most of the length of the plate. Subgenital plate elongated with moderately protruding triangular apex.</p>
            <p><bold><italic>Measurements</italic> (in mm).</bold><abbrev xlink:title="body length" id="ABBRID0EKFAE">BL</abbrev> (9.21); <abbrev xlink:title="pronotum length" id="ABBRID0EOFAE">PL</abbrev> (12.33); <abbrev xlink:title="pronotum width" id="ABBRID0ESFAE">PWS</abbrev> (2.19); <abbrev xlink:title="pronotum width" id="ABBRID0EWFAE">PWM</abbrev> (2.85); <abbrev xlink:title="pronotum height" id="ABBRID0E1FAE">PHM</abbrev> (2.38); <abbrev xlink:title="vertex width" id="ABBRID0E5FAE">VW</abbrev> (0.84); <abbrev xlink:title="compound eye width" id="ABBRID0ECGAE">CEW</abbrev> (0.52); <abbrev xlink:title="antennal groove width" id="ABBRID0EGGAE">AGW</abbrev> (0.23); <abbrev xlink:title="scutellum width" id="ABBRID0EKGAE">SW</abbrev> (0.3); <abbrev xlink:title="fore femur length" id="ABBRID0EOGAE">FFL</abbrev> (2.16); <abbrev xlink:title="fore femur width" id="ABBRID0ESGAE">FFW</abbrev> (0.45); <abbrev xlink:title="mid femur length" id="ABBRID0EWGAE">MFL</abbrev> (2.28); <abbrev xlink:title="mid femur width" id="ABBRID0E1GAE">MFW</abbrev> (0.49); <abbrev xlink:title="hind femur length" id="ABBRID0E5GAE">HFL</abbrev> (4.75); <abbrev xlink:title="hind femur width" id="ABBRID0ECHAE">HFW</abbrev> (1.58); <abbrev xlink:title="tegmen length" id="ABBRID0EGHAE">TL</abbrev> (1.47); <abbrev xlink:title="tegmen width" id="ABBRID0EKHAE">TW</abbrev> (0.58)</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Holotype" id="SECID0EOHAE">
            <title>Holotype.</title>
            <p>Adult female collected in early 1993 by Duffner in Ecuador, Sucumbíos Province on the upper Aguarico. The holotype is deposited at the SMNK (Karlsruhe, Germany). See Fig. <xref ref-type="fig" rid="F2">2</xref> and Suppl. material <xref ref-type="supplementary-material" rid="S1">1</xref>.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Exact label information" id="SECID0E3HAE">
            <title>Exact label information.</title>
            <p>“ECUADOR (Sucumbios) / Oberer Aguarico, Anfang / 1993 leg. Duffner // 101, // Staatl. Museum / für Naturkunde / Karlsruhe; / ex coll. Riede, / Inv. Nr. E-Orthop-2“</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="Terra typica" id="SECID0EBIAE">
            <title>Terra typica.</title>
            <p>Ecuador, Sucumbíos Province. No precise locality known.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="distribution" id="SECID0EGIAE">
            <title>Distribution.</title>
            <p>Known only from a single specimen.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0ELIAE">
            <title>Etymology.</title>
            <p>The new species is named after Prof. Dr. Matthias Reinschmidt, the current director of the Karlsruhe Zoo for his dedication and investments in nature protection in Ecuador and around the world.</p>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="arpha">C838E76D-D666-5688-A989-409370A77FD1</object-id>
              <label>Figure 2.</label>
              <caption>
                <p><italic><tp:taxon-name>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. <bold>A.</bold> Regions where <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> species have been found, red shapes represent type localities, star–<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic>, circle–<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic>, triangle–<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic>; <bold>B.</bold> Frontal view; <bold>C.</bold> Dorsal view; <bold>D.</bold> Ventral view; <bold>E.</bold> Abdominal apex in ventral view; <bold>F.</bold> Lateral view; <bold>G.</bold> Image of the 3D model we generated.</p>
              </caption>
              <graphic xlink:href="evolsyst-08-119-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_1056785.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/1056785</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
      <sec sec-type="﻿Identification key to the species of Phelene" id="SECID0EQIAE">
        <title>﻿Identification key to the species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></title>
        <table-wrap content-type="key" position="anchor" orientation="portrait">
          <table id="TID0EEOAC" rules="all">
            <tbody>
              <tr>
                <td rowspan="1" colspan="1">1</td>
                <td rowspan="1" colspan="1">Tall narrow pronotal crest above tegmina</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Low elongated pronotal crest</td>
                <td rowspan="1" colspan="1">2</td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">2</td>
                <td rowspan="1" colspan="1">Lower half of body distinctly black, upper half black or brown; pulvilli of hind tarsus triangular and sharp; alae dark with bright venation; female subgenital plate oval with slight tubercle at its apex</td>
                <td rowspan="1" colspan="1">
                  <italic>
                    <tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name>
                  </italic>
                </td>
              </tr>
              <tr>
                <td rowspan="1" colspan="1">–</td>
                <td rowspan="1" colspan="1">Body uniformly brown; pulvilli of hind tarsus rounded and blunt; alae and their venation dark brown; female subgenital plate oval with moderately protruding triangular apex</td>
                <td rowspan="1" colspan="1"><italic><tp:taxon-name>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</td>
              </tr>
            </tbody>
          </table>
        </table-wrap>
      </sec>
    </sec>
    <sec sec-type="﻿Discussion" id="SECID0EYLAE">
      <title>﻿Discussion</title>
      <sec sec-type="﻿General remarks" id="SECID0E3LAE">
        <title>﻿General remarks</title>
        <p>The new species is at first glance quite similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic>. The latter was until recently known only from a single male specimen (<xref ref-type="bibr" rid="B5">Cadena-Castañeda et al. 2021</xref>), but a recent examination of collections from French Guiana revealed a number of new individuals, of which many were female (<xref ref-type="bibr" rid="B15">Itrac-Bruneau et al. 2023</xref>). This, coupled with our own <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> specimens, allowed us to thoroughly compare our only female specimen with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> females and find some key differences, most notably in the shape of the subgenital plate. Furthermore, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. was collected in Ecuador, while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic> is known only from French Guiana. Interestingly, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> is known from both Peru and French Guiana, and is sometimes syntopical with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="maroon">maroon</tp:taxon-name-part></tp:taxon-name></italic>. The specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> reported by Itrac-Bruneau et al. (<xref ref-type="bibr" rid="B15">2023</xref>) differ from the specimen from Marcapata, Peru by the shape of the pronotal crest and may represent a separate species. However, our specimens collected in central Peru (Fig. <xref ref-type="fig" rid="F1">1</xref>) exhibit the same amount of variability in this trait as between the French Guiana and Marcapata specimens, indicating either that this trait can vary somewhat or that the distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> species is very complex. Unfortunately, photographs of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">P.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="turgida">turgida</tp:taxon-name-part></tp:taxon-name></italic> types are not available. It is still possible to describe tetrigid species based on clear morphological characters, but DNA studies are becoming increasingly necessary, especially to address interesting biogeographical observations.</p>
      </sec>
      <sec sec-type="﻿Phylogenetic hypotheses" id="SECID0EUPAE">
        <title>﻿Phylogenetic hypotheses</title>
        <p>Until now, the question of the position of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> in the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name> phylogeny remained completely unaddressed. The brief synonymization of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chiriquia">Chiriquia</tp:taxon-name-part></tp:taxon-name></italic> (<xref ref-type="bibr" rid="B4">Cadena-Castañeda and Cardona 2015</xref>) was never elaborated despite being interesting and potentially premonitory. These two genera are clearly not synonymous, but their similarities cannot simply be explained as convergence (<xref ref-type="bibr" rid="B5">Cadena-Castañeda et al. 2021</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chiriquia">Chiriquia</tp:taxon-name-part></tp:taxon-name></italic> are alike in their facial morphologies, lengths of first tarsal segments, the placement of pronotal crests, and the sideways projected lateral lobes, which could point to some degree of relatedness. The genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Otumba">Otumba</tp:taxon-name-part></tp:taxon-name></italic> Morse, 1900 seems to be related to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Chiriquia">Chiriquia</tp:taxon-name-part></tp:taxon-name></italic> and could provide further hints upon its urgently needed revision.</p>
        <p>Further, we have to consider the genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">Metrodora</tp:taxon-name-part></tp:taxon-name></italic> Bolívar, 1887. Some of its species, e.g., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="lutosa">lutosa</tp:taxon-name-part></tp:taxon-name></italic> Bolívar, 1887, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gibbinota">gibbinota</tp:taxon-name-part></tp:taxon-name></italic> (Bruner, 1910), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="panamae">panamae</tp:taxon-name-part></tp:taxon-name></italic> (Hebard, 1924) also exhibit the earlier listed similarities with <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part></tp:taxon-name></italic>, while <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="gibbinota">gibbinota</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Metrodora">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="panamae">panamae</tp:taxon-name-part></tp:taxon-name></italic> also have white-tipped antennae. This genus and by extension its tribe, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Metrodorini</tp:taxon-name-part></tp:taxon-name> Bolívar, 1887, is composed of likely related but ill-defined taxa, which is hindering further research (<xref ref-type="bibr" rid="B17">Kasalo et al. 2023</xref>). The relationship between <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Metrodorinae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name> should be closely examined and it may even find the latter to be synonymous with the former.</p>
        <p>Lastly, all these genera share the mentioned characters with many members of the predominantly African tribe <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Xerophyllini</tp:taxon-name-part></tp:taxon-name> Günther, 1979, currently attributed to the subfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Cladonotinae</tp:taxon-name-part></tp:taxon-name>. Therefore, the possibility that <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Lophotettiginae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Metrodorinae</tp:taxon-name-part></tp:taxon-name>, and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Xerophyllini</tp:taxon-name-part></tp:taxon-name> form a monophyletic group should be seriously considered and examined from both morphological and molecular angles. The morphology akin to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Morphopus">Morphopus</tp:taxon-name-part></tp:taxon-name></italic> (Bolívar, 1887) (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Xerophyllini</tp:taxon-name-part></tp:taxon-name>) could be basal in this group, which could explain all the different derived morphologies within it, as well as its distribution across the southern half of Gondwana. It should be noted that we do not consider Asian <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Metrodorinae</tp:taxon-name-part></tp:taxon-name> as true members of that subfamily and that all of the mentioned higher taxa require extensive revisions.</p>
      </sec>
      <sec sec-type="﻿Value of 3D data in taxonomy" id="SECID0E3VAE">
        <title>﻿Value of 3D data in taxonomy</title>
        <p>We here provide the first 3D data for any <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Tetrigidae</tp:taxon-name-part></tp:taxon-name>, though more are in preparation. Such 3D data may become an important resource in future taxonomic works, especially in concert with artificial intelligence and machine learning approaches. One of the large advantages of including 3D data in taxonomic descriptions is that virtual specimens will be readily available around the world, without the danger of sending valuable and irreplaceable holotypes and risking them getting lost or damaged. In addition, such virtual specimens are at least a minor backup in case a type gets lost. The 3D models are scaled to size and allow easy and accurate measurement of distances and also have the benefit of allowing for 3D geometric morphometric measurements. Modern high-throughput X-ray imaging methods already offer the possibility to digitize large numbers of insects in a short time (<xref ref-type="bibr" rid="B26">van de Kamp et al. 2018</xref>; <xref ref-type="bibr" rid="B23">Rühr et al. 2021</xref>). In the future, artificial intelligence will help to further automate data acquisition as well as large-scale data analysis (<xref ref-type="bibr" rid="B12">Heethoff and van de Kamp 2023</xref>). Even statistical species delimitation may become a viable approach potentially speeding up the process of taxonomy, an important improvement in a time of heavy diversity decline.</p>
      </sec>
    </sec>
    <sec sec-type="﻿Author contributions" id="SECID0EPWAE">
      <title>﻿Author contributions</title>
      <p>NK: first and last draft, taxonomy, measurements, figures; MH: last draft, specimen handling; TvdK: last draft, 3D digitalization; JS: last draft, taxonomy.</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>﻿Acknowledgements</title>
      <p>We thank Marcus Zuber and Elias Hamann for their assistance with the tomographic scans and Tomás Faragó for tomographic reconstruction. We thank Mathias Vielsäcker for imaging the specimens. We acknowledge the KIT Light Source for provision of instruments at their beamlines and thank the Institute for Beam Physics and Technology (IBPT) for the operation of the Karlsruhe Research Accelerator (KARA). We are grateful to Tomislav Domazet-Lošo for reading an early version of this manuscript and giving valuable suggestions. We acknowledge the help of Leibniz Institut zur Analyse des Biodiversitätswandels in publishing this article.</p>
    </ack>
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    <sec sec-type="supplementary-material">
      <title>Supplementary materials</title>
      <supplementary-material id="S1" position="float" orientation="portrait" xlink:type="simple">
        <object-id content-type="doi">10.3897/evolsyst.8.124285.suppl1</object-id>
        <object-id content-type="arpha">D4A20863-3086-541F-A3B8-139899CB8146</object-id>
        <label>﻿Supplementary material 1</label>
        <caption>
          <p>﻿3D reconstruction of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov.</p>
        </caption>
        <statement content-type="dataType">
          <label>Data type</label>
          <p>pdf</p>
        </statement>
        <statement content-type="notes">
          <label>Explanation note</label>
          <p>3D reconstruction of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Phelene">Phelene</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="reinschmidti">reinschmidti</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. Click on the image to start the interactive 3D view; use the menu to switch between the standard views.</p>
        </statement>
        <media xlink:href="evolsyst-08-119-s001.pdf" mimetype="application" mime-subtype="pdf" position="float" orientation="portrait" xlink:type="simple" id="oo_1056786.pdf">
          <uri content-type="original_file">https://binary.pensoft.net/file/1056786</uri>
        </media>
        <permissions>
          <license xlink:type="simple">
            <license-p>This dataset is made available under the Open Database License (http://opendatacommons.org/licenses/odbl/1.0). The Open Database License (ODbL) is a license agreement intended to allow users to freely share, modify, and use this Dataset while maintaining this same freedom for others, provided that the original source and author(s) are credited.</license-p>
          </license>
        </permissions>
        <attrib specific-use="authors">Thomas van de Kamp</attrib>
      </supplementary-material>
    </sec>
  </back>
</article>
