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  <front>
    <journal-meta>
      <journal-id journal-id-type="publisher-id">60</journal-id>
      <journal-id journal-id-type="index">urn:lsid:arphahub.com:pub:056820A7-C438-5162-B00B-FC18BD6C2AA0</journal-id>
      <journal-id journal-id-type="aggregator">urn:lsid:zoobank.org:pub:4B0FB9C5-4BE9-4A41-8BA6-2C2FD3522FC1</journal-id>
      <journal-title-group>
        <journal-title xml:lang="en">Zoologia</journal-title>
        <abbrev-journal-title xml:lang="en">Zoologia</abbrev-journal-title>
      </journal-title-group>
      <issn pub-type="epub">1984-4689</issn>
      <publisher>
        <publisher-name>Pensoft Publishers</publisher-name>
      </publisher>
    </journal-meta>
    <article-meta>
      <article-id pub-id-type="doi">10.3897/zoologia.34.e13230</article-id>
      <article-id pub-id-type="publisher-id">13230</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Animalia</subject>
          <subject>Gastropoda</subject>
          <subject>Stylommatophora</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Systematics</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic>, a new genus and new species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> from Brazil (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Gastropoda</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Stylommatophora</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</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>Gabriela Cuezzo</surname>
            <given-names>Maria</given-names>
          </name>
          <email xlink:type="simple">gcuezzo@webmail.unt.edu.ar</email>
          <uri content-type="orcid">https://orcid.org/0000-0002-3531-1744</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Pena</surname>
            <given-names>Meire Silva</given-names>
          </name>
          <xref ref-type="aff" rid="A2">2</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Instituto de Biodiversidad Neotropical, CONICET-UNT, Facultad de Ciencias Naturales, Universidad Nacional de Tucumán. Miguel Lillo 205, 4000 Tucumán, Argentina.</addr-line>
        <institution>Universidad Nacional de Tucumán</institution>
        <addr-line content-type="city">Tucumán</addr-line>
        <country>Argentina</country>
      </aff>
      <aff id="A2">
        <label>2</label>
        <addr-line content-type="verbatim">Laboratório de Malacologia, Museu de Ciências Naturais, Pontifícia Universidade Católica de Minas Gerais. Avenida Dom José Gaspar 500, Coração Eucarístico, Belo Horizonte, MG, Brazil.</addr-line>
        <institution>Pontifícia Universidade Católica de Minas Gerais</institution>
        <addr-line content-type="city">Belo Horizonte</addr-line>
        <country>Brazil</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Maria Gabriela Cuezzo (<email xlink:type="simple">gcuezzo@webmail.unt.edu.ar</email>)</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2017</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>27</day>
        <month>04</month>
        <year>2017</year>
      </pub-date>
      <volume>34</volume>
      <fpage>1</fpage>
      <lpage>12</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/8B68C230-32CE-59F6-99BE-6207BAE0F71C">8B68C230-32CE-59F6-99BE-6207BAE0F71C</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/1B7B6395-EE91-46AA-9774-89832FE0F47A">1B7B6395-EE91-46AA-9774-89832FE0F47A</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/576010">576010</uri>
      <history>
        <date date-type="received">
          <day>27</day>
          <month>05</month>
          <year>2016</year>
        </date>
        <date date-type="accepted">
          <day>29</day>
          <month>11</month>
          <year>2016</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Maria Gabriela Cuezzo, Meire Silva Pena</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/1B7B6395-EE91-46AA-9774-89832FE0F47A</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>We describe a new genus and a new species in the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name>, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. n.</bold> The new species was found at the National Park Cavernas do Peruaçu, in northern portion of the state of Minas Gerais, Brazil. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. n.</bold> is classified in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> based on the fact that it shares the following diagnostic features of the family: a dart apparatus with a single dart sac, and two unequal mucous glands at the terminal genitalia. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. n.</bold> differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> Doering, 1874 by having a granulose protoconch, shell spire with blunt apex, complex microsculpture on the teleoconch and closed umbilicus fused with the shell wall. Also, significant differences between the two genera are the presence of a long and thin kidney that extends more than half the length of the pulmonary cavity, the presence of a flagellar caecum, and a smooth jaw in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. n.</bold> The finding of this new species and genus is particularly significant to refine the definition of the family, since <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> has been traditionally diagnosed using the same characters of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic> Pilsbry &amp; Cockerell, 1937, the other valid genus in the family, is monospecific and is only known by the morphology of the shell. In many ways it is similar to <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>. A cladistics analysis was made in the present study which supports <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic><bold>gen. n.</bold> as a different entity and as sister group of the <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name>.</p>
      </abstract>
      <kwd-group>
        <label>Key words</label>
        <kwd>Cerrado</kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="family">Pleurodontidae</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="infraclass">Pulmonata</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>South America</kwd>
        <kwd>Taxonomy</kwd>
      </kwd-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="SECID0ENAAC">
      <title>Introduction</title>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> is a Pulmonate land snail family exclusively distributed in South America. It is composed of the genera <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> Doering, 1874 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic> Pilsbry &amp; Cockerell, 1937. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> is currently composed of 63 species distributed in Peru, Bolivia, Argentina, Paraguay and southern Brazil with a single extra occurrence in Colombia (<xref ref-type="bibr" rid="B8">Linares and Vega 2011</xref>). The species are classified into five subgenera (<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> s.s., <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Angrandiella">Angrandiella</tp:taxon-name-part></tp:taxon-name></italic> Ancey, 1886, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Doeringiana">Doeringiana</tp:taxon-name-part></tp:taxon-name></italic> Ihering, 1929, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Karlchmidtia">Karlchmidtia</tp:taxon-name-part></tp:taxon-name></italic> Hass, 1955 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pilsbrya">Pilsbrya</tp:taxon-name-part></tp:taxon-name></italic> Ancey, 1887) (<xref ref-type="bibr" rid="B23">Zilch 1959</xref>, <xref ref-type="bibr" rid="B17">Richardson 1982</xref>, <xref ref-type="bibr" rid="B5">Cuezzo 2006</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic> is a monotypic genus, known only by <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">D.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="harringtoni">harringtoni</tp:taxon-name-part></tp:taxon-name></italic> Pilsbry &amp; Cockerel, 1937 from Bolivia.</p>
      <p><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> is currently diagnosed by the same synapomorphies of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> because <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic> is only known by its original description, which is entirely based on shell characters. Based on a cladistic hypothesis (<xref ref-type="bibr" rid="B5">Cuezzo 2006</xref>), <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> is characterized by the following synapomorphies i) malleated shell body whorl surface with diagonal ribs, ii) umbilicus overlapping, but not fused to the body whorl, iii) thick, widely reflexed peristome, iv) presence of a dart sac apparatus inserting in the vagina, or directly into the atrium, v) mucous glands unequal in size and shape, vi) insertion of mucous glands ducts in middle portion of dart sac, and vii) penial retractor muscle inserting in epiphallus medial portion. The short duct of the bursa copulatrix, a character that traditionally had been used to define the genus, is characteristic only of a small group of species.</p>
      <!--PageBreak-->
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> was classified by <xref ref-type="bibr" rid="B13">Pilsbry (1894)</xref> in the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Helicidae</tp:taxon-name-part></tp:taxon-name>, tribe Belogona Euadenia. The members of the latter are distinguished by having mucous glands of typically glandular structure, in contrast to the tube-like glands of the Belogona Siphonadenia. Thiele (1929) later classified <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Fruticicolidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Epiphragmophorinae</tp:taxon-name-part></tp:taxon-name>. <xref ref-type="bibr" rid="B14">Pilsbry (1939)</xref> assembled all the American dart-bearing helices in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Helminthoglyptidae</tp:taxon-name-part></tp:taxon-name> and restricted <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> to the South American <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Epiphragmophorinae</tp:taxon-name-part></tp:taxon-name> Hoffman, 1928 from the same family. <xref ref-type="bibr" rid="B10">Nordsieck (1987)</xref> maintained <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Epiphragmophorinae</tp:taxon-name-part></tp:taxon-name> Hoffman, 1928, but moving it into <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Xanthonychidae</tp:taxon-name-part></tp:taxon-name>, while stating that its reproductive system resembles that of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Cepoliinae</tp:taxon-name-part></tp:taxon-name>: diverticulum usually missing, one dart sac, dart glands unequal, one elongate and the other compact, inserting on the dart sac or on its base. Finally, <xref ref-type="bibr" rid="B19">Schileyko (1991)</xref> elevated <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Epiphragmophorinae</tp:taxon-name-part></tp:taxon-name> to family within <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</tp:taxon-name-part></tp:taxon-name> and this classification was followed by <xref ref-type="bibr" rid="B1">Bouchet and Rocroi (2005)</xref> in the last gastropod family nomenclator. <xref ref-type="bibr" rid="B5">Cuezzo (2006)</xref> studied species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> from Argentina and part of Bolivia, providing the first phylogenetic analysis of the genus. Species from Peru and Paraguay have been scarcely investigated and are mostly known by their original descriptions. Specimens with their entire body preserved are rare in malacological collections globally. Consequently, comprehensive anatomical or molecular studies are not feasible in most cases. In Brazil there is a single species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="oresigena">oresigena</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="subspecies" reg="bernardius">bernardius</tp:taxon-name-part></tp:taxon-name></italic> Ihering in Pilsbry, 1900, known from Serra da Bocaina and Campos do Jordão in São Paulo state and from state of Rio Grande do Sul.</p>
      <p>During a field trip to the National Park Cavernas do Peruaçu in northern Minas Gerais, Brazil, to collect gastropods, a striking group of land snails was found. Analysis of the specimens collected revealed that they represent a new species of the family <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name>. The objective of the present work is to describe the new species in a new genus of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> and to discuss its position among the South American <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</tp:taxon-name-part></tp:taxon-name>.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EXIAC">
      <title>Material and methods</title>
      <p>Snails were collected at the National Park Cavernas do Peruaçu (<named-content content-type="dwc:verbatimCoordinates"><named-content content-type="geo-json" specific-use="{&quot;type&quot;:&quot;Point&quot;,&quot;coordinates&quot;:[-44.600000,-14.933333]}" id="NCID0EAJAC">14°56’S, 44°36’W</named-content></named-content>) located in the state of Minas Gerais, Brazil. This conservation unit was created in 1999 with 143,353.84 ha (<ext-link xlink:type="simple" ext-link-type="uri" xlink:href="http://www.icmbio.gov.br">http://www.icmbio.gov.br</ext-link>), to protect limestone caverns. The calcareous massif is covered by rare and typical deciduous (Caatinga) or semi-deciduous forests called Seasonal Dry Tropical Forest (SDTF) (<xref ref-type="bibr" rid="B12">Pennington et al. 2000</xref>, <xref ref-type="bibr" rid="B16">Prado 2000</xref>). Open shrub savannah (Cerrado) also occurs on the top of the mountains, and in poorly drained areas of the Park where the Peruaçu River originates. Shrub density is lower at the watersheds, and there the vegetation is totally herbaceous (Campo Limpo and Veredas), with localized Buriti palm-trees. The Caatinga, the Cerrado and the Chaco dry areas are extensive, open biomes, and form the dry diagonal in South America, which is a natural phytogeographic unit (<xref ref-type="bibr" rid="B12">Pennington et al. 2000</xref>, <xref ref-type="bibr" rid="B16">Prado 2000</xref>). The Cerrado spreads across 2,031,990 km<sup>2</sup> of the central Brazilian Plateau and is the second largest of Brazil’s major biomes, after the Amazon. This biodiversity hotspot actually receives abundant rainfall (between 1,100 and 1,600 mm per year), although this rainfall is concentrated in a six to seven month period between October and April. The rest of the year is characterized by a pronounced dry season.</p>
      <p>Live specimens and dry shells of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. were collected from rocky outcrops in dry deciduous forests of the National Park. Dry shells adhered to rocks or to leaves were abundant but live snails were scarce. The collected specimens were drowned in water for relaxation previous to fixation in ethanol 96%. Their shells were then photographed and measured using the software ImageJ 1.49 (Fig. <xref ref-type="fig" rid="F1">1</xref>). Anatomical information was obtained by dissecting specimens and studying them under a Leica MZ6 stereoscope, illustrations of the dissected parts where made with the aid of a camera lucida. Photographs of the different organ systems were taken using a Nikon 5000 camera attached to the stereoscope. They were enhanced and finalized using the software Corel Draw version X3. The terminology for the anatomical descriptions follows <xref ref-type="bibr" rid="B21">Tompa (1984)</xref>. The terms proximal and distal refer to the position of an organ or part of an organ in relation to the gamete flow from the ovotestis (proximal) to the genital pore (distal), as in previous studies (<xref ref-type="bibr" rid="B3">Cuezzo 1997</xref>, <xref ref-type="bibr" rid="B5">Cuezzo 2006</xref>). The limit between the epiphallus and penis is based on the internal sculpture of their inner walls. The radula and the jaw of specimens were observed and photographed with a Jeol Scanning Electron Microscopy 35CF at the Integral Center of Electron Microscopy of the National University of Tucumán, Argentina. Shell microphotographs were obtained with a DSM 950ZEISS SEM at “Centro de Aquisição e Processamento de Imagens” of the Federal University of Minas Gerais, Brazil.</p>
      <p>Intitutional abbreviations used in the text: IBN, Instituto de Biodiversidad Neotropical, Tucumán, Argentina; MLP-Ma, Museo de La Plata, Buenos Aires, Argentina; MNRJ, Museu Nacional Rio de Janeiro, Universidade Federal do Rio de Janeiro, Brazil; MCN, Museu Ciências Naturais, Pontifícia Universidade de Minas Gerais, Belo Horizonte, Brazil.</p>
      <p>For the cladistic analysis, a matrix of 35 characters from the general anatomy plus shell morphology was generated for 24 terminal taxa (Appendix 1), following characters and codifications of <xref ref-type="bibr" rid="B4">Cuezzo (2003)</xref> for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part></tp:taxon-name></italic> Fischer, 1807 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Labyrinthus">Labyrinthus</tp:taxon-name-part></tp:taxon-name></italic> Beck, 1837 and <xref ref-type="bibr" rid="B5">Cuezzo (2006)</xref> for species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>, with modifications. The characters used in the analysis are listed in Appendix 2. In the text, characters and state characters numbers are located between parenthesis. Only species for which the anatomy had been described were included in the character matrix. The data matrix was built in Winclada, v. 1.00.08 (<xref ref-type="bibr" rid="B9">Nixon 2002</xref>). Non-applicable data were coded as “–” and multistate characters were treated as additive. Cladistic analyses were performed with TNT, version 1.5 (<xref ref-type="bibr" rid="B7">Goloboff et al. 2008</xref>) with Character Weighting and Traditional Search basing <!--PageBreak-->the strategy on RAS + TBR (random addition sequences plus swap by tree bisection and reconnection), with 1,000 replications and 100 trees saved per replication. The default concavity (K) value was used in all analyses (K = 3000). In the analysis, trees were rooted in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part></tp:taxon-name></italic> based on <xref ref-type="bibr" rid="B22">Wade et al. (2007)</xref> hypothesis on the phylogenetic relationships of the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</tp:taxon-name-part></tp:taxon-name>. Clade support was estimated using symmetric resampling (<xref ref-type="bibr" rid="B6">Goloboff et al. 2003</xref>), because the resulting values obtained under this procedure are not distorted by character weighting. Additionally, support for the obtained clades was calculated using the Jackknife resampling method.</p>
      <fig id="F1" position="float" orientation="portrait">
        <object-id content-type="doi">10.3897/zoologia.34.e13230.figure1</object-id>
        <object-id content-type="zenodo_dep_id">993967</object-id>
        <object-id content-type="arpha">DA943370-1037-5A24-97A3-257268EF93F3</object-id>
        <label>Figure 1.</label>
        <caption>
          <p>Shell measurements. Abbreviations (AD) dorsal view area, (Aap) apertural area, (Abw) body whorl area, (AL) lateral view area, (Dap) apertural diameter, (DM) major diameter, (Dm) minor diameter, (H) total shell height, (Hap) height of aperture, (Hbw) body whorl height, (Pbw) body whorl perimeter, (PD) dorsal view shell perimeter, (PL) lateral view shell perimeter, (PS) parietal space.</p>
        </caption>
        <graphic xlink:href="zoologia-34-e13230-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133816.jpg">
          <uri content-type="original_file">https://binary.pensoft.net/fig/133816</uri>
        </graphic>
      </fig>
    </sec>
    <sec sec-type="TAXONOMY" id="SECID0EWNAC">
      <title>Taxonomy</title>
      <p>Supra superfamily classification follows <xref ref-type="bibr" rid="B1">Bouchet and Rocroi (2005)</xref> and <xref ref-type="bibr" rid="B15">Ponder and Lindberg (1997)</xref>. This classification tries to integrate the results of recent cladistics work by using the unranked “clade” above the rank of superfamily while still using the traditional Linnaean ranks for superfamilies and all taxa below the rank of superfamily.</p>
      <sec sec-type="Class Gasteropoda Cuvier, 1795" id="SECID0EEOAC">
        <title>Class <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Gasteropoda</tp:taxon-name-part></tp:taxon-name> Cuvier, 1795</title>
        <sec sec-type="Heterobranchia" id="sec1">
          <title>Clade <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superorder">Heterobranchia</tp:taxon-name-part></tp:taxon-name> Burmeister, 1837</title>
          <sec sec-type="Stylommatophora" id="sec2">
            <title>Clade <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Stylommatophora</tp:taxon-name-part></tp:taxon-name> Schmidt, 1855</title>
            <sec sec-type="Helicoidea" id="sec3">
              <title>Superfamily <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</tp:taxon-name-part></tp:taxon-name> Rafinesque, 1815</title>
              <sec sec-type="Epiphragmophoridae" id="sec4">
                <title><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> Hoffmann, 1928</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">Gasteropoda</named-content>
                      </kwd>
                      <kwd>
                        <named-content content-type="family" xlink:type="simple">Epiphragmophoridae</named-content>
                      </kwd>
                    </kwd-group>
                  </tp:treatment-meta>
                  <tp:nomenclature>
                    <tp:taxon-name><object-id content-type="arpha">78462639-7B66-5C6B-B162-32A4A0A2EC4D</object-id>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part>
                      <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/48537C28-29CA-4488-97FF-1C570990D9F9</object-id>
                    </tp:taxon-name>
                    <tp:taxon-status>gen. n.</tp:taxon-status>
                  </tp:nomenclature>
                  <tp:treatment-sec sec-type="diagnosis" id="SECID0EOAAE">
                    <title>Diagnosis.</title>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> gen. n. is distinguished by the following characters: 1) shell globose with blunt apex; 2) protoconch sculptured with granules; 3) teleoconch sculptured with complex microstructures; 4) umbilicus imperforate, parietal wall fused with columellar zone of peristome; 5) wavy spiral lines below the periphery and over ventral teleoconch surface; 6) genitalia with a dart apparatus composed by a single dart sac and two unequal mucous glands, one globose and the other oval; 7) presence of a flagellar caecum; 8) bursa copulatrix duct short, no longer than the sac.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="type species" id="SECID0E2AAE">
                    <title>Type species.</title>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. by original designation.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="description" id="SECID0EMBAE">
                    <title>Description.</title>
                    <p>Shell globose, with 4 to 5 convex whorls. Spire conic with blunt apex. Protoconch granulose. Teleoconch sculptured. Wavy spiral grooves at the ventral teleoconch surface. Aperture subcircular with thin peristome. Umbilicus closed. Presence of spiral brownish bands more pronounced in the body whorl. Kidney long and thin, more than half the lung roof length. Genitalia with a dart apparatus and two unequal mucous glands.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="etymology" id="SECID0ERBAE">
                    <title>Etymology.</title>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> is a compound name formed by <italic>Minas</italic> in honor to the Brazilian state where the species was found, and <italic>selates</italic>, a noun in the genitive singular, that derives from the Greek meaning “snail” (<xref ref-type="bibr" rid="B2">Brown 1979</xref>).</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="remarks" id="SECID0ELCAE">
                    <title>Remarks.</title>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> gen. n. is classified in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> because it has a dart apparatus and two unequal mucous glands at the terminal genitalia. These structures are diagnostic of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Helicoidea</tp:taxon-name-part></tp:taxon-name>) and their morphology serve to differentiate this family from the remaining helicoidean groups. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic> differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> in its depressed shell with an acute peripheral keel and open umbilicus. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> differs from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> in its general shell shape with blunt apex, granulose protoconch and complex sculpture of the teleoconch surface. The wavy spiral grooves at the ventral teleoconch surface in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> are lacking in both, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Dinotropis">Dinotropis</tp:taxon-name-part></tp:taxon-name></italic>. The presence of a long and thin kidney in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> is very different to the kidney shape in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>, which is triangular and shorter.</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>
                        <named-content content-type="order" xlink:type="simple">Gasteropoda</named-content>
                      </kwd>
                      <kwd>
                        <named-content content-type="family" xlink:type="simple">Epiphragmophoridae</named-content>
                      </kwd>
                    </kwd-group>
                  </tp:treatment-meta>
                  <tp:nomenclature>
                    <tp:taxon-name><object-id content-type="arpha">8BA4AD32-FDCE-5F87-BBFC-3F8FE38E0783</object-id>
                      <tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part>
                      <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part>
                      <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/9AACED00-8AFB-4736-85DC-14A3399A04CB</object-id>
                    </tp:taxon-name>
                    <tp:taxon-status>sp. n.</tp:taxon-status>
                    <xref ref-type="fig" rid="F1">Figs 1</xref>
                    <xref ref-type="fig" rid="F2">, 2–7</xref>
                    <xref ref-type="fig" rid="F3">, 8–13</xref>
                    <xref ref-type="fig" rid="F4">, 14–19</xref>
                    <xref ref-type="fig" rid="F5">, 20–21</xref>
                    <xref ref-type="fig" rid="F6">, 22–26</xref>
                  </tp:nomenclature>
                  <tp:treatment-sec sec-type="diagnosis" id="SECID0EGHAE">
                    <title>Diagnosis.</title>
                    <p>Shell globular, with three spiral continuous pigmented bands, the middle, equatorial band thinner. Protoconch <!--PageBreak-->granulose. Dorsal side of teleoconch with axial lines bearing triangular lamellae, ventral teleoconch with wavy, concentric, spiral grooves. Imperforate umbilicus. Jaw smooth. Kidney triangular, long and thin, of about 60 to 70% the length of the lung roof. Vas deferens insertion in lower portion of flagellar caecum. Strong, short muscular penial retractor inserting at proximal epiphallus.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="etymology" id="SECID0EOHAE">
                    <title>Etymology.</title>
                    <p>The species name derives from the Greek <italic>paradoxos</italic> meaning “strange, contrary to expectation” (<xref ref-type="bibr" rid="B2">Brown 1979</xref>) as this is a species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name> that was not expected to occur in the state of Minas Gerais, Brazil.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="description" id="SECID0EFIAE">
                    <title>Description.</title>
                    <p>Shell (Figs <xref ref-type="fig" rid="F2">2</xref>–<xref ref-type="fig" rid="F3">13</xref>) dextral, helicoidal with 4¾ convex, solid whorls. Coloration pale brown with three spiral darker brown bands more separated from each other in the body whorl. Medial pigmented band thinner than the other bands (Figs <xref ref-type="fig" rid="F2">2–7</xref>). Suture impressed. Protoconch of 1½ whorls, covered by oval, tightly arranged pustules (Figs <xref ref-type="fig" rid="F3">8</xref>, <xref ref-type="fig" rid="F3">9</xref>). Dorsal side of teleoconch with slightly curved axial growth lines that, at higher magnifications, appear as axial thin lines bearing broad-base triangular lamellae. These axial lines are not continuous and are separated by narrow spaces with wrinkles. Each lamellae has an axis from the base to the upper extreme of the triangle (Figs <xref ref-type="fig" rid="F3">10–13</xref>). The long axes of the lamellae are perpendicular to the shell sutures. This sculpture looks like granules at naked eye. Wavy spiral grooves below the periphery, and in the basal surface area. Peristome thin, reflexed, some specimens showing a basal thickening and or an incipient palatal tooth. Aperture roundish, without angulation, and rounded in the palatal zone, well reflected in the columellar zone covering the umbilicus. Shell imperforate. Shell measurements. Table <xref ref-type="table" rid="T1">1</xref>, Fig. <xref ref-type="fig" rid="F1">1</xref>. Digestive system. Radula (Figs <xref ref-type="fig" rid="F4">14–17</xref>) Central tooth long, unicuspid. Lateral teeth long, with incipient lateral cusps, of about 54–56 µm (n = 10). Marginal teeth bicuspid, similar to laterals in shape and size. Jaw (Figs <xref ref-type="fig" rid="F4">18</xref>, <xref ref-type="fig" rid="F4">19</xref>) Horseshoe slightly arched, translucent, with no division. Surface almost smooth with thin, transverse grooves visible with high magnification. Pallial system (Figs <xref ref-type="fig" rid="F5">20</xref>, <xref ref-type="fig" rid="F5">21</xref>) Kidney triangular, long and thin, of about 60 to 70% the length of the lung roof. Main pulmonary vein thick, splitting into two secondary branches before reaching mantle collar; pulmonary roof dark grey in color, furrowed by well marked, but lesser minor transverse veins. Secondary ureter runs parallel to rectum, completely closed until reaching mantle collar. Ureteric interramus triangular in shape, deeply excavated. Genital System (Figs <xref ref-type="fig" rid="F6">22–26</xref>) Terminal genitalia with dart apparatus and two mucous glands (Figs <xref ref-type="fig" rid="F6">22–24</xref>). Right ommatophore retractor crosses the distal genital system between penis and vagina. Vagina long. Penis and vagina entering side by side in atrium. Bursa copulatrix with a globular sac and short, thick duct. Bursa copulatrix slightly longer than free oviduct. Single dart sac muscular, with medial constriction, ending in the atrium. Upper dart sac inverted pear shaped, lower portion of dart sac bellow constriction cylindrical, thicker (Fig. <xref ref-type="fig" rid="F6">24</xref>). Two unequal mucous glands with their respective thin efferent ducts inserting independently above the dart sac constriction (Fig. <xref ref-type="fig" rid="F6">25</xref>). One of the glands bean shaped with medial duct and the other oval with one end bulkier than the other. Vas deferens is a long, narrow duct that passes between one of the mucous glands and the dart sac, over the penial sheath, going down it adheres to penis-vagina at angle with connective tissue and then rises parallel to the penis complex to insert bellow the flagellar caecum. Vas deferens insertion marks the limit between epiphallus and flagellum. Penial complex tubular without external differentiation between penis and epiphallus. Short distal muscular penis sheath. Penial retractor muscle thick inserting in proximal epiphallus. Penial cavity occupied by several thin wavy pilasters and an oval verge sculptured with overlapping lamellae (Fig. <xref ref-type="fig" rid="F6">26</xref>). Penis longer than epiphallus, limits differentiated through their particular inner sculpture. Penial sheath short of about 1/5 of penial length. Epiphallus inner cavity with three thick pilasters. Proximal epiphallus with a rounded caecum where the vas deferens inserts. Flagellum tubular, longer than epiphallus. Spermoviduct long with uterus zone plicated transversal to the longitudinal axis. Albumen gland bean shaped with a prominent fertilization pouch-spermathecal complex (FPSC). Atrium short.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="type locality" id="SECID0ETKAE">
                    <title>Type locality.</title>
                    <p>Brazil, <italic>Minas Gerais</italic>: Itacarambi, National Park Cavernas do Peruaçu, Vale dos Sonhos (523m, X = 0599645, Y = 8343426), M.S. Pena, A. Suhett, D.C. Souza leg., December 2010, (MNRJ 34.580), Holotype (ethanol preserved specimen).</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="material" id="SECID0EBLAE">
                    <title>Other material examined.</title>
                    <p>Brazil, <italic>Minas Gerais</italic>: Itacarambi, National Park Cavernas do Peruaçu, Nossa Senhora Aparecida Farm (532 m, X = 0589284, Y = 8328970), M.S. Pena, A. Suhett, <!--PageBreak--><!--PageBreak--><!--PageBreak-->D.C. Souza leg., December 2010, (IBN 21-S, MLP-Ma 14216), dry shells, (IBN 861), ethanol preserved specimens. Paratypes. Brazil, <italic>Minas Gerais</italic>: Itacarambi, National Park Cavernas do Peruaçu, Brejal, Peruaçu River side (663 m, X = 0579404, Y = 8332170), (MCN 192), dry shells. Brazil, <italic>Minas Gerais</italic>: Itacarambi, Natiomal Park Cavernas do Peruaçu, Janelão Cave (714 m, X = 0581514, Y = 8329046) M.S. Pena, A. Suhett, D.C. Souza leg., December 2010, (MCN 208).</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="distribution" id="SECID0EDMAE">
                    <title>Distribution.</title>
                    <p>Thus far known only from National Park Cavernas do Peruaçu, northern region of Minas Gerais, Brazil.</p>
                  </tp:treatment-sec>
                  <tp:treatment-sec sec-type="remarks" id="SECID0EPMAE">
                    <title>Remarks.</title>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> differs from all known species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> by having a granulose protoconch, the shell spire with a blunt apex, and by the wavy, spiral grooves in the ventral region of the shell. The fused, imperforate umbilicus on the shells of this new species is not typical of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora.</tp:taxon-name-part></tp:taxon-name></italic> Most of species of the genus present an open, perspective umbilicus. The exceptions are <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="argentina">argentina</tp:taxon-name-part></tp:taxon-name></italic> (Holmberg, 1909) and some specimens of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="variegata">variegata</tp:taxon-name-part></tp:taxon-name></italic> Hylton Scott, 1962. The presence of wavy spiral grooves at the base of the shell of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. is noteworthy, sharply contrasting with the condition found in all other species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>, where it is absent, except for <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> (<tp:taxon-name-part taxon-name-part-type="subgenus" reg="Pylsbrya">Pylsbrya</tp:taxon-name-part>) <tp:taxon-name-part taxon-name-part-type="species" reg="farrisi">farrisi</tp:taxon-name-part></tp:taxon-name> (Pfeiffer, 1859), which has shallow spiral lines. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. also differs from the species classified in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> in the shape and length of the kidney, and the smooth jaw. In the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> for which the anatomy has been studied, the kidney is no more than half the length of the pulmonary roof, while the jaw is ribbed. A noteworthy character present in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. is the presence of a flagellar caecum, a structure not found in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>. In this new species, the vas deferens inserts in the lower portion of the caecum. The penial retractor muscle in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> is mostly long and thin, inserting in the epiphallus, while in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. this muscle is stronger, inserting in the caecum. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. is isolated from the area of distribution of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>, whose area of highest species richness is in the western portion of South America.</p>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. resembles some species of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pleurodontidae</tp:taxon-name-part></tp:taxon-name> by the presence of complex structures in the terminal genitalia of the male, such as the flagellar caecum. It is also similar to some <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pleurodontidae</tp:taxon-name-part></tp:taxon-name> in its long and thin kidney, the crowded granules in the surface of the shell protoconch, the globular general shape of the shell, the complex microsculpture on the teleoconch and in its smooth jaw. The presence of a flagellar caecum is noteworthy in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic>, this structure being absent in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Epiphragmophoridae</tp:taxon-name-part></tp:taxon-name>, while it is characteristic <!--PageBreak--><!--PageBreak-->of some <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pleurodontidae</tp:taxon-name-part></tp:taxon-name> such as <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Polydontes">Polydontes</tp:taxon-name-part></tp:taxon-name></italic> Montfort, 1810 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="incerta">incerta</tp:taxon-name-part></tp:taxon-name></italic> (Férussac, 1823) (<xref ref-type="bibr" rid="B4">Cuezzo 2003</xref>).</p>
                    <p>In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic>, however, the vas deferens inserts in the flagellar pouch while in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Polydontes">Polydontes</tp:taxon-name-part></tp:taxon-name></italic> the vas deferens inserts above this caecum. At first glance, the shell of this new species is very similar to some species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Pleurodontidae</tp:taxon-name-part></tp:taxon-name>), except for the presence of concentric sculpture below the periphery, and in the basal area. It also differs in the morphology of the terminal genitalia that has a dart complex and a different flagellum shape. In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part></tp:taxon-name></italic>, the vas deferens is twisted around the epiphallus, descending to the peni-oviducal angle, while in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. the vas deferens runs straight, parallel to the penis complex, without looping around the epiphallus. Most of the Helicoidean families have a ribbed jaw or ‘odontognath jaw’, except for the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Sagdidae</tp:taxon-name-part></tp:taxon-name> with a ‘stegognath jaw’ and the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Sphicterochilidae</tp:taxon-name-part></tp:taxon-name> and <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Cepoliinae</tp:taxon-name-part></tp:taxon-name> with a smooth jaw (‘oxygnath jaw’) (<xref ref-type="bibr" rid="B4">Cuezzo 2003</xref>). In <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">M.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. the jaw was found to be smooth, only having fine transverse lines visible with electron microscopy, similar to jaws in <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Caracolus">Caracolus</tp:taxon-name-part></tp:taxon-name></italic> Montfort, 1810 and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Labyrinthus">Labyrinthus</tp:taxon-name-part></tp:taxon-name></italic>.</p>
                    <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n. is distributed within a National Park area where Cerrado and Caatinga are the dominant biomes. These are considered high diversity hotspot areas. Within these hotspots areas specimens were collected in typical deciduous or semi-deciduous forests called Seasonally Dry Tropical Forests (SDTF) (<xref ref-type="bibr" rid="B12">Pennington et al. 2000</xref>, <xref ref-type="bibr" rid="B16">Prado 2000</xref>). These types of forest are drought-adapted, tree-dominated ecosystems with a more or less continuous canopy. Grasses are not a significant element in these forests, which are currently scattered in eastern South America. A hypothesis has been advanced predicting that SDTFs were more widespread during drier glacial climates, and that their fragmentation during the current wet interglacial period has contributed to speciation and the distribution patters of species observed today (<xref ref-type="bibr" rid="B12">Pennington et al. 2000</xref>, <xref ref-type="bibr" rid="B16">Prado 2000</xref>). This hypothesis is for the most part based on the assessment of floristic links among species assemblages in the SDTF areas (<xref ref-type="bibr" rid="B18">Sarkinen et al. 2011</xref>). Land snails inhabiting this type of dry forest, from the Cerrado and Caatinga hotspots areas, have been scarcely studied and consequently their conservation status is unknown. Most vertebrates and plants have been more extensively documented in the Cerrado and Caatinga hotspots areas (<xref ref-type="bibr" rid="B11">Overbeck et al. 2015</xref>), but this is not the case with many invertebrate groups.</p>
                    <table-wrap id="T1" position="float" orientation="portrait">
                      <label>Table 1.</label>
                      <caption>
                        <p>Shell dimensions in mm or mm<sup>2</sup> (n = 14). DM major diameter, Dm minor diameter, AD dorsal area, PD dorsal perimenter, H total height, HBw body whorl height, AL lateral area, PL lateral perimeter, ABw body whorl area, PBw body whorl perimeter, Dap diameter of the aperture, Hap height of the aperture, EP length parietal space, AAp apertural area, Pap apertural perimeter, DP penultimate whorl diameter, DPr protoconch diameter (see Fig. <xref ref-type="fig" rid="F1">1</xref>).</p>
                      </caption>
                      <table id="TID0EAYAE" rules="all">
                        <tbody>
                          <tr>
                            <th rowspan="1" colspan="1">Character</th>
                            <th rowspan="1" colspan="1">Mean</th>
                            <th rowspan="1" colspan="1">SD</th>
                            <th rowspan="1" colspan="1">Min</th>
                            <th rowspan="1" colspan="1">Max</th>
                            <th rowspan="1" colspan="1">Holotype</th>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">DM</td>
                            <td rowspan="1" colspan="1">28.526</td>
                            <td rowspan="1" colspan="1">0.765</td>
                            <td rowspan="1" colspan="1">27.103</td>
                            <td rowspan="1" colspan="1">29.714</td>
                            <td rowspan="1" colspan="1">28.934</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">Dm</td>
                            <td rowspan="1" colspan="1">25.582</td>
                            <td rowspan="1" colspan="1">0.669</td>
                            <td rowspan="1" colspan="1">24.182</td>
                            <td rowspan="1" colspan="1">26.374</td>
                            <td rowspan="1" colspan="1">26.058</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">AD</td>
                            <td rowspan="1" colspan="1">557.304</td>
                            <td rowspan="1" colspan="1">30.211</td>
                            <td rowspan="1" colspan="1">502.938</td>
                            <td rowspan="1" colspan="1">596.790</td>
                            <td rowspan="1" colspan="1">566.908</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">PD</td>
                            <td rowspan="1" colspan="1">85.289</td>
                            <td rowspan="1" colspan="1">2.375</td>
                            <td rowspan="1" colspan="1">80.699</td>
                            <td rowspan="1" colspan="1">88.531</td>
                            <td rowspan="1" colspan="1">86.135</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">H</td>
                            <td rowspan="1" colspan="1">18.663</td>
                            <td rowspan="1" colspan="1">1.005</td>
                            <td rowspan="1" colspan="1">17.183</td>
                            <td rowspan="1" colspan="1">20.370</td>
                            <td rowspan="1" colspan="1">19.375</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">HBw</td>
                            <td rowspan="1" colspan="1">15.068</td>
                            <td rowspan="1" colspan="1">0.878</td>
                            <td rowspan="1" colspan="1">13.904</td>
                            <td rowspan="1" colspan="1">16.351</td>
                            <td rowspan="1" colspan="1">15.904</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">AL</td>
                            <td rowspan="1" colspan="1">375.374</td>
                            <td rowspan="1" colspan="1">30.818</td>
                            <td rowspan="1" colspan="1">324.118</td>
                            <td rowspan="1" colspan="1">411.659</td>
                            <td rowspan="1" colspan="1">386.807</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">PL</td>
                            <td rowspan="1" colspan="1">74.999</td>
                            <td rowspan="1" colspan="1">3.084</td>
                            <td rowspan="1" colspan="1">69.055</td>
                            <td rowspan="1" colspan="1">78.120</td>
                            <td rowspan="1" colspan="1">76.548</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">ABw</td>
                            <td rowspan="1" colspan="1">343.840</td>
                            <td rowspan="1" colspan="1">28.940</td>
                            <td rowspan="1" colspan="1">299.433</td>
                            <td rowspan="1" colspan="1">379.724</td>
                            <td rowspan="1" colspan="1">355.669</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">PBw</td>
                            <td rowspan="1" colspan="1">74.081</td>
                            <td rowspan="1" colspan="1">2.727</td>
                            <td rowspan="1" colspan="1">69.154</td>
                            <td rowspan="1" colspan="1">77.544</td>
                            <td rowspan="1" colspan="1">74.445</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">Dap</td>
                            <td rowspan="1" colspan="1">14.886</td>
                            <td rowspan="1" colspan="1">1.091</td>
                            <td rowspan="1" colspan="1">12.997</td>
                            <td rowspan="1" colspan="1">16.888</td>
                            <td rowspan="1" colspan="1">15.004</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">Hap</td>
                            <td rowspan="1" colspan="1">14.937</td>
                            <td rowspan="1" colspan="1">0.598</td>
                            <td rowspan="1" colspan="1">13.647</td>
                            <td rowspan="1" colspan="1">15.763</td>
                            <td rowspan="1" colspan="1">15.158</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">PS</td>
                            <td rowspan="1" colspan="1">11.382</td>
                            <td rowspan="1" colspan="1">0.610</td>
                            <td rowspan="1" colspan="1">10.245</td>
                            <td rowspan="1" colspan="1">12.335</td>
                            <td rowspan="1" colspan="1">11.158</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">AAp</td>
                            <td rowspan="1" colspan="1">190.094</td>
                            <td rowspan="1" colspan="1">14.988</td>
                            <td rowspan="1" colspan="1">158.663</td>
                            <td rowspan="1" colspan="1">208.360</td>
                            <td rowspan="1" colspan="1">208.360</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">PAp</td>
                            <td rowspan="1" colspan="1">52.650</td>
                            <td rowspan="1" colspan="1">2.173</td>
                            <td rowspan="1" colspan="1">49.201</td>
                            <td rowspan="1" colspan="1">56.690</td>
                            <td rowspan="1" colspan="1">53.222</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">DP</td>
                            <td rowspan="1" colspan="1">16.605</td>
                            <td rowspan="1" colspan="1">0.874</td>
                            <td rowspan="1" colspan="1">14.900</td>
                            <td rowspan="1" colspan="1">17.782</td>
                            <td rowspan="1" colspan="1">17.782</td>
                          </tr>
                          <tr>
                            <td rowspan="1" colspan="1">DPr</td>
                            <td rowspan="1" colspan="1">3.050</td>
                            <td rowspan="1" colspan="1">0.264</td>
                            <td rowspan="1" colspan="1">2.651</td>
                            <td rowspan="1" colspan="1">3.550</td>
                            <td rowspan="1" colspan="1">2.873</td>
                          </tr>
                        </tbody>
                      </table>
                    </table-wrap>
                    <fig id="F2" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figures2-7</object-id>
                      <object-id content-type="zenodo_dep_id">993969</object-id>
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                      <label>Figures 2–7.</label>
                      <caption>
                        <p>Shell morphology: (2) Dorsal view of Holotype, MNRJ 34.580; (3) lateral position of holotype shell and soft body; (4) dorsal, (5) ventral, and (6) lateral view of paratype, IBN 861; (7) live snail. Scale bar: 3–5 = 10 mm.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133817.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133817</uri>
                      </graphic>
                    </fig>
                    <fig id="F3" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figures8-13</object-id>
                      <object-id content-type="zenodo_dep_id">993971</object-id>
                      <object-id content-type="arpha">0CF37CA2-F3BB-55C0-B3FE-AA641C807800</object-id>
                      <label>Figures 8–13.</label>
                      <caption>
                        <p>Shell morphology: (8) Protoconch in dorsal view, scale bar = 4mm; (9) lateral view of the protoconch and first whorls, scale bar = 4mm; (10) general view of the teleoconch microsculpture, scale bar = 100μ; (11) body whorl microsculpture consisting on axial rows of triangular lamella separated by wrinkles; (12) perpendicular view of the lamella, scale bar = 5μ; (13) detail of a triangular lamellae showing its central axis, scale bar = 2 μm.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g003.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133818.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133818</uri>
                      </graphic>
                    </fig>
                    <fig id="F4" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figures14-19</object-id>
                      <object-id content-type="zenodo_dep_id">993973</object-id>
                      <object-id content-type="arpha">17DFF5F0-4D3E-5B47-954E-890EA38272F9</object-id>
                      <label>Figures 14–19.</label>
                      <caption>
                        <p>Digestive system: (14) Dorso-lateral view of the radula, arrows points to central tooth in two transverse rows of teeth; (15) detail on a dorso-lateral view of the lateral teeth;(16) detail of lateral teeth close to margin in dorsal view (17) margin of the radula showing curve shaped marginal teeth; (18) dorsal view of the smooth, fragile jaw; (19) detail of the dorsal surface of the jaw showing transversal shallow grooves. Scale bars = 10 μm.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g004.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133819.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133819</uri>
                      </graphic>
                    </fig>
                    <fig id="F5" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figures20-21</object-id>
                      <object-id content-type="zenodo_dep_id">993975</object-id>
                      <object-id content-type="arpha">BCFA5F6B-45E4-5173-8DBE-5BEDDF073877</object-id>
                      <label>Figures 20–21.</label>
                      <caption>
                        <p>Pallial system: (20) photograph of the ventral zone of the pulmonary roof, note the long kidney, with respect to the total length of the lung; (21) line drawing of the same region detailing the limits between kidney and ureter and the shallow veins crossing the pulmonary roof. Scale bar: 21 = 5mm. Abbreviations (k) kidney, (mc) mantle collar, (pv) pericardic vein, (r) rectum, (su) secondary ureter.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g005.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133820.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133820</uri>
                      </graphic>
                    </fig>
                    <fig id="F6" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figures22-26</object-id>
                      <object-id content-type="zenodo_dep_id">993977</object-id>
                      <object-id content-type="arpha">1745383F-082B-530D-9891-F4DBEC0EA5AA</object-id>
                      <label>Figures 22–26.</label>
                      <caption>
                        <p>Genital system: (22, 23) line drawing and photograph of the complete dissected out reproductive system (ag) albumen gland, (bc) bursa copulatrix, (ds) dart sac, (ec) flagellar caecum, (fl) flagellum, (go) genital opening, (hd) hermaphroditic duct, (mg1) mucous gland 1, (mg2) mucous gland 2, (p) penis, (pr) penial retractor muscle, (s) spermoviduct, (vd) vas deferens; (24) detail of the terminal genitalia (pl) penial plates, (ps) penial sheath, (v) verge; (25) vas deferens and insertion of mucous gland ducts into dart sac (mgd) mucous gland duct; (26) inner sculpture of the penial complex showing the verge inside the penis. Scale bars: 22 and 25 = 5 mm.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g006.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133821.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133821</uri>
                      </graphic>
                    </fig>
                    <fig id="F7" position="float" orientation="portrait">
                      <object-id content-type="doi">10.3897/zoologia.34.e13230.figure27</object-id>
                      <object-id content-type="zenodo_dep_id">993979</object-id>
                      <object-id content-type="arpha">484909E6-B060-5707-9003-79C430ED819B</object-id>
                      <label>Figure 27.</label>
                      <caption>
                        <p>Consensus tree obtained from two trees under implied weight approach highlighting the position of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> gen. n. Only taxa with described anatomy were included in the matrix. Numbers below branches are node numbers. Symmetric resampling values (left number) and jackknife values (right number) are located above branches. Only values above 50% are illustrated.</p>
                      </caption>
                      <graphic xlink:href="zoologia-34-e13230-g007.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_133822.jpg">
                        <uri content-type="original_file">https://binary.pensoft.net/fig/133822</uri>
                      </graphic>
                    </fig>
                  </tp:treatment-sec>
                </tp:taxon-treatment>
              </sec>
            </sec>
          </sec>
        </sec>
      </sec>
    </sec>
    <sec sec-type="CLADISTIC ANALYSIS" id="SECID0EFFAG">
      <title>Cladistic analysis</title>
      <p>The main goal of the analysis performed was to evaluate and support the description of a new genus. For this, only the species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> with known complete anatomical and shell morphology information were used. The morphological data set analyzed under the implied weights approach resulted in two most parsimonious trees. The resulting strict consensus tree is illustrated (Fig. <xref ref-type="fig" rid="F7">27</xref>). Synapomorphies that support <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> gen. n. as a new genus, different from <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> are: body whorl surface covered with triangular lamellae <!--PageBreak-->(0: 5); duct of bursa copulatrix extremely short, not longer than sac (22:0); presence of a flagellar caecum (32:1).</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> (node 29) was also recovered as a monophyletic genus in all optimal trees, supported by the following synapomorphies: Body whorl surface with axial ribs regularly distributed (0:3); umbilicus overlapped but not solded to body whorl (1: 1); protoconch smooth (8:0), and Penial retractor muscle inserting in medial zone of epiphallus (21:1). Within <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic>, the clade [<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="variegata">variegata</tp:taxon-name-part></tp:taxon-name></italic> [<italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="hemiclausa">hemiclausa</tp:taxon-name-part></tp:taxon-name></italic> [<italic>E. tucumanensis+<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="argentina">argentina</tp:taxon-name-part></tp:taxon-name></italic>]]] (node 27) has the highest SR and jackknife support and resulted monophyletic in all trees obtained.</p>
      <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part></tp:taxon-name></italic> gen. n. is the sister group of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> in both optimal trees. This relationship is supported by the following synapomorphies: Presence of mucous glands in terminal genitalia (10: 1); presence of a dart apparatus (11:1); medium to short length of the bursa copulatrix duct (22:01) and a finger-like short to medium flagellum (25:1).</p>
    </sec>
  </body>
  <back>
    <ack>
      <title>Acknowledgements</title>
      <p>Thanks to the Instituto Chico Mendes de Conservação da Biodiversidade for the working permits and support (SISBIO-ICMBIO 19133-1 de 08/04/2009) and Fundo de Incentivo à Pesquisa (FIP), Pontifícia Universidade Católica de Minas Gerais for financial support to MSP. MGC is a researcher of the Argentine National Council for Scientific Research (CONICET). Financial support to MGC has been received through PIP 0055 (CONICET). We also thank the anonymous reviewers and the editor for their valuable comments.</p>
    </ack>
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      <app id="S1">
        <title>Appendix 1</title>
        <sec sec-type="Appendix 1" id="sec5">
          <title/>
          <table-wrap id="T2" position="float" orientation="portrait">
            <label/>
            <caption>
              <p>Character matrix used in cladistic analysis. Only taxa with described anatomy were included in the matrix. Character codification according to <xref ref-type="bibr" rid="B4">Cuezzo (2003</xref>, <xref ref-type="bibr" rid="B5">2006</xref>) with some modifications. “–”are non-applicable characters, “?” are missing characters.</p>
            </caption>
            <table id="TID0EOLAG" rules="all">
              <tbody>
                <tr>
                  <td rowspan="2" colspan="1"/>
                  <td rowspan="1" colspan="1"/>
                  <td rowspan="1" colspan="34">Characters</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1"/>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">4</td>
                  <td rowspan="1" colspan="1">5</td>
                  <td rowspan="1" colspan="1">6</td>
                  <td rowspan="1" colspan="1">7</td>
                  <td rowspan="1" colspan="1">8</td>
                  <td rowspan="1" colspan="1">9</td>
                  <td rowspan="1" colspan="1">10</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">4</td>
                  <td rowspan="1" colspan="1">5</td>
                  <td rowspan="1" colspan="1">6</td>
                  <td rowspan="1" colspan="1">7</td>
                  <td rowspan="1" colspan="1">8</td>
                  <td rowspan="1" colspan="1">9</td>
                  <td rowspan="1" colspan="1">20</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">4</td>
                  <td rowspan="1" colspan="1">5</td>
                  <td rowspan="1" colspan="1">6</td>
                  <td rowspan="1" colspan="1">7</td>
                  <td rowspan="1" colspan="1">8</td>
                  <td rowspan="1" colspan="1">9</td>
                  <td rowspan="1" colspan="1">30</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">4</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">4</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">–</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="argentina">argentina</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">[03]</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="cryptomphala">cryptomphala</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">[12]</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="escoipensis">escoipensis</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="hemiclausa">hemiclausa</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="guevarai">guevarai</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="hieronymi">hieronymi</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="jujuyensis">jujuyensis</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="parodizi">parodizi</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="puella">puella</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">4</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">?</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="puntana">puntana</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="quirogai">quirogai</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="rhathymos">rhathymos</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="saltana">saltana</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">3</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="tomsici">tomsici</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="trenquelleonis">trenquelleonis</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">[01]</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
                <tr>
                  <td rowspan="1" colspan="1">
                    <italic>
                      <tp:taxon-name>
                        <tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">E.</tp:taxon-name-part>
                        <tp:taxon-name-part taxon-name-part-type="species" reg="trifasciata">trifasciata</tp:taxon-name-part>
                      </tp:taxon-name>
                    </italic>
                  </td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">2</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">1</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                  <td rowspan="1" colspan="1">0</td>
                </tr>
              </tbody>
            </table>
          </table-wrap>
        </sec>
      </app>
      <app id="S2">
        <title>Appendix 2</title>
        <sec sec-type="Character list used in the cladistics analysis of species" id="SECID0EMFBI">
          <title/>
          <p>
      Character list used in the cladistics analysis of species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Epiphragmophora">Epiphragmophora</tp:taxon-name-part></tp:taxon-name></italic> with known anatomy plus the new genus and species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Minaselates">Minaselates</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="paradoxa">paradoxa</tp:taxon-name-part></tp:taxon-name></italic> sp. n.</p>
          <p>0. Body whorl surface: with thin growth lines = 0; with thick growth ridges = 1; malleated with diagonal ribs = 2; with axial ribs regularly distributed = 3; pustules/granules to wrinkles = 4; triangular lamella in axial rows = 5. [additive].</p>
          <p>1. Umbilicus: Fused with basal lip of peristome = 0; overlapped but not fused to body whorl = 1; perspective wide not overlapped by peristomal lip = 2; perspective narrow slightly overlapped = 3; wide partly overlapped = 4.</p>
          <p>2. Shape of the aperture: sub circular = 0; oval horizontal = 1; sub quadrangular = 2.</p>
          <p>3. Peristome: Thin expanded slightly reflexed = 0; thick wide reflexed = 1; thin highly expanded = 2.</p>
          <p>4. Basal callus in peristome: absent = 0; present = 1.</p>
          <p>5. Peripheral bands: absent = 0; present = 1.</p>
          <p>6. Body whorl periphery: convex = 0; equatorially subcarinated = 1; supraequatorially subcarinated = 2; carinated = 3. [additive].</p>
          <p>7. Aperture respect to body whorl: not descending = 0; descending = 1.</p>
          <p>8. Protoconch sculpture: smooth = 0; granulose = 1.</p>
          <p>9. Spire apex: pointed = 0; dull = 1; not evident = 2.</p>
          <p>10. Mucous glands in terminal genitalia: absent = 0; present = 1.</p>
          <p>11. Dart apparatus: absent = 0; present = 1.</p>
          <p>12. Shape of dart sac: long finger-like usually with constriction = 0; short, cylindrical, no constriction = 1.</p>
          <p>13. Dart sac insertion: in vagina = 0; in atrium = 1.</p>
          <p>14. Dart sac papillae: absent = 0; present = 1.</p>
          <p>15. Relation between ducts of both mucous glands: separated = 0; distally fused or contiguous = 1.</p>
          <p>16. Position of left mucous gland duct: distal respect to the body of the gland = 0; equatorial respect to the body of the gland = 1.</p>
          <p>17. Shape of right mucous gland: not sac-like = 0; sac-Like = 1.</p>
          <p>18. Right mucous gland: not fused with atrium wall = 0; distally fused with atrium wall = 1.</p>
          <p>19. Penis length respect to epiphallus: half epiphallus length = 0; as long as epiphallus = 1; longer than epiphallus length = 2.</p>
          <p>20. Penial papillae (= verge): absent = 0; present = 1.</p>
          <p>21. Penial retractor muscle: inserts in distal epiphallus = 0; inserts in medial zone of epiphallus = 1; inserts in proximal epiphallus = 2; inserts in proximal penis = 3.</p>
          <p>22. Duct of bursa copulatrix: extremely short not longer than sac = 0; medium = 1; long = 2. [additive].</p>
          <p>23. Vagina: short = 0; medium to long = 1; extremely long = 2. [additive].</p>
          <p>24. Atrium:short = 0; medium to long = 1.</p>
          <p>25. Flagellum: thin, long = 0; finger-like, short to medium = 1; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pleurodonte">Pleurodonte</tp:taxon-name-part></tp:taxon-name></italic>-like = 2; <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Labyrinthus">Labyrinthus</tp:taxon-name-part></tp:taxon-name></italic>-like = 3. [additive].</p>
          <p>26. Penial muscular band: absent = 0; present = 1.</p>
          <p>27. Penial sheath (penial tunica): simple = 0; double or multi<!--PageBreak-->layer = 1.</p>
          <p>28. Microhabitat associated to: rocks = 0; tree trunks = 1.</p>
          <p>29. Vas deferens: surrounding dart sac = 0; not surrounding dart sac = 1.</p>
          <p>30. Epiphallus proximal portion: Not widen at point entrance vas deferens = 0; Widen at point of entrance of vas deferens = 1.</p>
          <p>31. Penial retractor: not = 0; forming a loop around vas deferens before insertion in epiphallus = 1.</p>
          <p>32. Flagellar caecum: absent = 0; present = 1.</p>
          <p>33. Kidney length respect to pulmonary roof: not exceeding half of pulmonary roof length = 0; more than half the pulmonary roof = 1.</p>
          <p>34. Jaw: ribbed = 0; smooth = 1.</p>
        </sec>
      </app>
    </app-group>
  </back>
</article>
