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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.36.e36293</article-id>
      <article-id pub-id-type="publisher-id">36293</article-id>
      <article-categories>
        <subj-group subj-group-type="heading">
          <subject>Research Article</subject>
        </subj-group>
        <subj-group subj-group-type="biological_taxon">
          <subject>Crabronidae</subject>
        </subj-group>
        <subj-group subj-group-type="scientific_subject">
          <subject>Systematics</subject>
        </subj-group>
      </article-categories>
      <title-group>
        <article-title>A new fossil species of the crabronid wasp genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name>) from Dominican amber</article-title>
      </title-group>
      <contrib-group content-type="authors">
        <contrib contrib-type="author" corresp="yes">
          <name name-style="western">
            <surname>Melo</surname>
            <given-names>Gabriel A.R.</given-names>
          </name>
          <email xlink:type="simple">garmelo@ufpr.br</email>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
        <contrib contrib-type="author" corresp="no">
          <name name-style="western">
            <surname>Rosa</surname>
            <given-names>Brunno B.</given-names>
          </name>
          <uri content-type="orcid">https://orcid.org/0000-0002-7342-9612</uri>
          <xref ref-type="aff" rid="A1">1</xref>
        </contrib>
      </contrib-group>
      <aff id="A1">
        <label>1</label>
        <addr-line content-type="verbatim">Laboratório de Biologia Comparada de Hymenoptera, Departamento de Zoologia, Universidade Federal do Paraná. Caixa Postal 19020, 81530-980 Curitiba, Brazil.</addr-line>
        <institution>Universidade Federal do Paraná</institution>
        <addr-line content-type="city">Curitiba</addr-line>
        <country>Brazil</country>
      </aff>
      <author-notes>
        <fn fn-type="corresp">
          <p>Corresponding author: Gabriel A.R. Melo (<email xlink:type="simple">garmelo@ufpr.br</email>)</p>
        </fn>
        <fn fn-type="edited-by">
          <p>Editorial responsibility: Gabriel L.F. Mejdalani</p>
        </fn>
      </author-notes>
      <pub-date pub-type="collection">
        <year>2019</year>
      </pub-date>
      <pub-date pub-type="epub">
        <day>01</day>
        <month>11</month>
        <year>2019</year>
      </pub-date>
      <volume>36</volume>
      <fpage>1</fpage>
      <lpage>5</lpage>
      <uri content-type="arpha" xlink:href="http://openbiodiv.net/8CB6E709-7BE3-5872-95B3-31339C805F57">8CB6E709-7BE3-5872-95B3-31339C805F57</uri>
      <uri content-type="zoobank" xlink:href="http://zoobank.org/70D21FB8-3942-4D80-8AF7-1D6F2B1CAC3D">70D21FB8-3942-4D80-8AF7-1D6F2B1CAC3D</uri>
      <uri content-type="zenodo_dep_id" xlink:href="https://zenodo.org/record/3533493">3533493</uri>
      <history>
        <date date-type="received">
          <day>17</day>
          <month>05</month>
          <year>2019</year>
        </date>
        <date date-type="accepted">
          <day>28</day>
          <month>08</month>
          <year>2019</year>
        </date>
      </history>
      <permissions>
        <copyright-statement>Gabriel A.R. Melo, Brunno B. Rosa</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/70D21FB8-3942-4D80-8AF7-1D6F2B1CAC3D</self-uri>
      <abstract>
        <label>Abstract</label>
        <p>The first fossil species of the wasp genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> from Dominican amber is described. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic><bold>sp. nov.</bold> is based on a single female specimen exhibiting a somewhat generalized morphology compared to the extant fauna. While the new species resembles the Neotropical species in a few characters, it does not possess the many specialized features exhibited by this species group, especially those related to the female legs. This is the 4<sup>th</sup> genus and the 7<sup>th</sup> species of apoid wasps described from Dominican amber.</p>
      </abstract>
      <kwd-group>
        <label>Key words</label>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="superfamily">Apoidea</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>
          <tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part>
          </tp:taxon-name>
        </kwd>
        <kwd>fossil</kwd>
        <!--PageBreak-->
        <kwd>taxonomy</kwd>
      </kwd-group>
      <funding-group>
        <award-group>
          <funding-source>
            <named-content content-type="funder_name">Conselho Nacional de Desenvolvimento Científico e Tecnológico</named-content>
            <named-content content-type="funder_identifier">501100003593</named-content>
            <named-content content-type="funder_doi">http://doi.org/10.13039/501100003593</named-content>
          </funding-source>
        </award-group>
      </funding-group>
    </article-meta>
  </front>
  <body>
    <sec sec-type="Introduction" id="SECID0EVE">
      <title>Introduction</title>
      <p>The amber from the Dominican Republic is an important fossiliferous deposit because of its abundance and availability, fine preservation, in addition to rare kinds of inclusions (<xref ref-type="bibr" rid="B6">Grimaldi and Engel 2005</xref>, <xref ref-type="bibr" rid="B9">Penney 2010</xref>). The detailed reconstruction of the ecosystem indicates a vanished tropical forest (<xref ref-type="bibr" rid="B4">Grimaldi 1995</xref>, <xref ref-type="bibr" rid="B5">1996</xref>). The age has been estimated to be Early Miocene through early Middle Miocene, about 15 to 20 million years ago, according to available biostratigraphic and paleogeographic data (<xref ref-type="bibr" rid="B7">Iturralde-Vinent and MacPhee 1996</xref>).</p>
      <p>More than 200 species of hymenopterans have been described from Dominican amber inclusions and only six of them are apoid wasps (<xref ref-type="bibr" rid="B9">Penney 2010</xref>). They all belong to <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Crabroninae</tp:taxon-name-part></tp:taxon-name> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name>): three species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">Trypoxylon</tp:taxon-name-part></tp:taxon-name></italic> Latreille, 1796, two of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Pison">Pison</tp:taxon-name-part></tp:taxon-name></italic> Jurine, 1808 (both genera in <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Trypoxylini</tp:taxon-name-part></tp:taxon-name>), and one of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Lindenius">Lindenius</tp:taxon-name-part></tp:taxon-name></italic> Lepeletier, 1839 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part></tp:taxon-name>) (<xref ref-type="bibr" rid="B10">Pulawski 2019</xref>). The <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part></tp:taxon-name> comprises the largest tribe of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name> with a world sum of 48 genera and 1,498 species (<xref ref-type="bibr" rid="B10">Pulawski 2019</xref>). <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> Morawitz, 1866 has currently 17 species, eight of them in the Palearctic region, three in the Nearctic, and six in the Neotropical region (<xref ref-type="bibr" rid="B8">Melo and Rosa 2015</xref>). Additionally, the genus has three fossil species, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="mortuellus">mortuellus</tp:taxon-name-part></tp:taxon-name></italic> Cockerell, 1906 from Florissant, Colorado (Miocene), and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="succinalis">succinalis</tp:taxon-name-part></tp:taxon-name></italic> (Cockerell, 1909) and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="tornquisti">tornquisti</tp:taxon-name-part></tp:taxon-name></italic> (Cockerell, 1909) from Baltic amber (<xref ref-type="bibr" rid="B10">Pulawski 2019</xref>). In this paper we describe the first fossil species of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> from Dominican amber.</p>
    </sec>
    <sec sec-type="materials|methods" id="SECID0EEBAC">
      <title>Material and methods</title>
      <p>The studied material belongs to the Amber Fossil Collection, Division of Invertebrate Zoology, American Museum of Natural History (AMNH). Terminology for external morphology follows <xref ref-type="bibr" rid="B2">Bohart and Menke (1976)</xref> with a few modifications. The photographs were obtained with a LEICA DFC295 digital camera attached to the stereoscopic microscope LEICA M125, and finished in the software Zerene Stacker (1.04 version build).</p>
    </sec>
    <sec sec-type="Taxonomy" id="SECID0EOBAC">
      <title>Taxonomy</title>
      <sec sec-type="Hymenoptera Linnaeus Apoidea Latreille Crabronidae Latreille Tracheliodes Morawitz" id="SECID0ESBAC">
        <title><tp:taxon-name>
            <tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part>
          </tp:taxon-name> Linnaeus <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="superfamily">Apoidea</tp:taxon-name-part></tp:taxon-name> Latreille <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name> Latreille <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> Morawitz</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">Hymenoptera</named-content>
              </kwd>
              <kwd>
                <named-content content-type="family" xlink:type="simple">Crabronidae</named-content>
              </kwd>
            </kwd-group>
          </tp:treatment-meta>
          <tp:nomenclature>
            <tp:taxon-name><object-id content-type="arpha">166F4003-3807-57A8-87E3-8CA255EA9ABE</object-id>
              <tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part>
              <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part>
              <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/CB5DF3D6-3B55-4115-8AC4-693B70FA4F21</object-id>
            </tp:taxon-name>
            <tp:taxon-status>sp. nov.</tp:taxon-status>
            <xref ref-type="fig" rid="F1">Figs 1–4</xref>
            <xref ref-type="fig" rid="F2">, 5–9</xref>
          </tp:nomenclature>
          <tp:treatment-sec sec-type="diagnosis" id="SECID0E6DAC">
            <title>Diagnosis.</title>
            <p><italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. can be set apart from the living taxa by the following combination of characters: (1) Thick setae along lower margin of clypeus relatively short, their length subequal to diameter of antennal socket; (2) Clypeal disc weakly convex, without medial protuberance; (3) Distance between antennal socket and eye orbit short, subequal to the distance between sockets; (4) Upper frons and vertex relatively flat, without conspicuous undulations of the integument; (5) Fore femur fusiform, not expanded ventrally, its lower surface devoid of thick bristles; (6) Mid femur fusiform, not expanded dorsally; (7) Mid tibial spur unmodified and relatively short; (8) Hind femur strongly thickened in the basal third, its anterior surface mostly flat; (9) First metasomal segment petiolate, dorsal surface of petiole convex; (10) Pygidial plate tear-shaped, with relatively weak preapical narrowing.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="description" id="SECID0EQEAC">
            <title>Description.</title>
            <p>Female holotype. <bold>Measurements</bold> (in mm). Approximate body length, 6.5; maximum head width, 1.9. <bold>Pilosity.</bold> Head and mesosoma mostly glabrous with sparse, fine and very short appressed hairs. Lower margin of clypeus with preapical line of long setae, their length subequal to diameter of antennal socket, lateral ones longer, thicker and slightly curved inwards; clypeal disc with sparser and shorter setae on its upper half; lower margin of gena with short and dense pilosity; parocular area, along scapal basin, with a dense strip of short, decumbent hairs facing outwards; dorsal surface of scape with numerous decumbent micro-setae. Anterior half of lateral portion of mesepisternum, behind omaular carina, with conspicuously dense short pilosity. Foreleg: lower surface of femur devoid of thick bristles; basitarsus with a row of short and thick bristles along inner margin, its posterior margin with a row of five bristles, bristles almost as long as diameter of basitarsus. Midleg: femur with thick bristles on posterior surface, inner surface with erect bristles directed towards apex, those on ventral surface slightly longer; anterior surface of tibia, right above the spur, with a pair of conspicuous lanceolate bristles; apical third of tibia with short thick setae on outer surface, these similar to those on remainder of tibia (not forming spininiform bristles), except for an apical pair of thick spiniform bristles. Hindleg: trochanter with a conspicuous patch of short decumbent setae; femur with thick relatively long bristles on anterior surface, bristles erect and directed towards apex; tibia with sparse, very thick, short bristles on outer surface; basitarsus with row of thick and short bristles forming a comb along its inner margin. Metasoma without conspicuous pilosity, except for few short setae on ventral portion of S6 and on margin of T6 bordering pygidial plate. <bold>Integumental surface. <!--PageBreak--></bold>Head and mesosoma finely microreticulated, dull. Pronotum with few parallel carinae in front of pronotal lobe. Posterior two-thirds of lateral surface of mesepisternum, metepisternum, and lateral portion of propodeum finely microstriated, striation more conspicuous on mesepisternum and metepisternum; omaular carina well developed and reaching mesepisternal sulcus ventrally, adjacent region behind carina with small foveae; mesepisternal sulcus weakly indicated and not foveolated. <bold>Structure.</bold> Head: cly<!--PageBreak-->peus transverse, its lateral portions very narrow, its lower margin with three broad teeth in the middle, lateral teeth aligned with antennal socket, disc weakly convex without projection; tentorial pit situated at the transition between inner and lower orbits, area around it weakly depressed; distance between antennal sockets subequal to the distance from socket to the inner orbit of compound eyes; scapal basin relatively shallow, only slightly depressed relative to lateral surface; distance between lower inner orbits approximately 0.7× width of eye in frontal view; gena in lateral view approximately 1.2× as wide as compound eye; upper frons and vertex relatively flat, without conspicuous undulations of the integument; ocello-orbital distance 1.6× the distance between posterior ocelli; distance between anterior and posterior ocelli about half the distance between the posterior ocelli; facial fovea linear, its length about 1.4﻿ the distance between posterior ocelli. Foreleg: femur fusiform, not expanded ventrally. Midleg: femur fusiform, not expanded dorsally, its lower portion flat, apical portion of anterior surface weakly excavated ventrally; tibial spur short, less than one-third of basitarsal length; basitarsus cylindrical, oval to circular in section. Hindleg: trochanter basically circular in section; femur strongly thickened in the basal third, its maximum height approximately 2.5× apical height, anterior surface mostly flat; hind tibia clavate as in other species of the genus; basitarsus gently curved and approximately 1.35× as long as mid basitarsus; inner tibial spur about 1.8× as long as outer spur. Metasoma: 1st segment petiolate and gradually broadening posteriorly; width of petiole approximately one-fourth of apical width of T1, in lateral view; dorsal surface of petiole convex; pygidial plate tear-shaped, flat, with relatively weak preapical narrowing, its margins limited by a low carina.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="type material" id="SECID0ECFAC">
            <title>Type material.</title>
            <p>Holotype female, in amber piece AMNH no. PB- 118, bearing the following labels: ‘Amber: Dominican Republic,\Specific Locality unknown\(Oligomiocene). Purchased in\Santo Domingo by Paul F. Burke.\AMNH no. PB- 118\Identifications:/<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Sphecidae</tp:taxon-name-part></tp:taxon-name>/<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Platypodidae</tp:taxon-name-part></tp:taxon-name>’ ‘PB118’. The piece measures 12 x 10 x 6 mm, and possesses a dark reddish orange color. The amber forms a darker reddish halo around the wasp specimen. The wasp wings are folded downward, making it difficult to observe both the wings and the metasoma; also, there are numerous air bubbles around its metasoma. There is a small beetle (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Platypodinae</tp:taxon-name-part></tp:taxon-name>, <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Curculionidae</tp:taxon-name-part></tp:taxon-name>) at ca. 2 mm above the holotype.</p>
          </tp:treatment-sec>
          <tp:treatment-sec sec-type="etymology" id="SECID0E3FAC">
            <title>Etymology.</title>
            <p>The species is named after the curator of AMNH’s amber collection, David A. Grimaldi, who has kindly made the material available for study.</p>
            <fig id="F1" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zoologia.36.e36293.figures1-4</object-id>
              <object-id content-type="zenodo_dep_id">3533495</object-id>
              <object-id content-type="arpha">67A2DE3E-7923-59E8-80D8-4D2C26BD1E33</object-id>
              <label>Figures 1–4.</label>
              <caption>
                <p>Holotype female of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (1) Head, frontal view; (2) Head and anterior portion of mesosoma, dorsal view; (3) Habitus, lateral view; (4) Habitus, ventral view. Scale bars = 1mm (Figs <xref ref-type="fig" rid="F1">1</xref> and <xref ref-type="fig" rid="F1">2</xref>, and Figs <xref ref-type="fig" rid="F1">3</xref> and <xref ref-type="fig" rid="F1">4</xref>, respectively at same scale).</p>
              </caption>
              <graphic xlink:href="zoologia-36-e36293-g001.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_352428.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/352428</uri>
              </graphic>
            </fig>
            <fig id="F2" position="float" orientation="portrait">
              <object-id content-type="doi">10.3897/zoologia.36.e36293.figures5-9</object-id>
              <object-id content-type="zenodo_dep_id">3533497</object-id>
              <object-id content-type="arpha">A60C632E-0CFB-5698-BEE3-908CA298CF4D</object-id>
              <label>Figures 5–9.</label>
              <caption>
                <p>Holotype female of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. (5) Head, ventral view; (6) Head and anterior portion of mesosoma, lateral view; (7) Posterior portion of mesosoma and anterior portion of metasoma, lateral view; (8) Mid and hind right legs, lateroventral view; Fig. <xref ref-type="fig" rid="F2">9</xref>. Pygidial plate, dorsal view. Scale bars = 1mm (Figs <xref ref-type="fig" rid="F2">6</xref> and <xref ref-type="fig" rid="F2">7</xref>, at same scale).</p>
              </caption>
              <graphic xlink:href="zoologia-36-e36293-g002.jpg" position="float" orientation="portrait" xlink:type="simple" id="oo_352429.jpg">
                <uri content-type="original_file">https://binary.pensoft.net/fig/352429</uri>
              </graphic>
            </fig>
          </tp:treatment-sec>
        </tp:taxon-treatment>
      </sec>
    </sec>
    <sec sec-type="Discussion" id="SECID0EBGAC">
      <title>Discussion</title>
      <p>Its worldwide distribution, with few and morphologically divergent species, and relatively ancient fossil record (Baltic amber) suggests that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> represents a relictual lineage within the <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part></tp:taxon-name>. Indeed, in <xref ref-type="bibr" rid="B1">Bennett’s (2010)</xref> phylogenetic study of the tribe, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> came out in a relatively isolated position, at the base of his <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Rhopalum">Rhopalum</tp:taxon-name-part></tp:taxon-name></italic> series. The genus differs from other crabronine genera by its elongate fore trochanter and a broad frons with the antennal sockets well separated from the inner orbits, which are only moderately convergent below (<xref ref-type="bibr" rid="B2">Bohart and Menke 1976</xref>). Also, <xref ref-type="bibr" rid="B1">Bennett (2010)</xref> lists as putative synapomorphies for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> the loss of the silver setal patch of the clypeus, elongate F1, elongate fore trochanter, an anteromedially flat T1, lack of lateral articular line on T2, and additional features of the male terminalia. As far as it is known, all species prey on ants (references in <xref ref-type="bibr" rid="B8">Melo and Rosa 2015</xref>).</p>
      <p>Considering the composition of the Dominican amber fauna (<xref ref-type="bibr" rid="B9">Penney 2010</xref>), one would expect that <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. should have a closer relationship to the species of the extant Neotropical fauna. This new species, however, has a somewhat generalized morphology and does not show a particular resemblance to any of the extant Neotropical species. The Neotropical species group is very characteristic and shares numerous unique conditions: (1) Lateral lobes of clypeus with very long setae; (2) Anterior tentorial pit situated in deep fovea; (3) Ventral surface of fore femur strongly expanded and with a row of specialized short bristles; (4) Dorsal surface of mid femur strongly expanded; (5) Bristles on apex of outer surface of mid tibia arranged in two parallel rows; (6) Metasomal petiole well defined, at least a long as one-third of the length of T1, with its dorsal surface completely flat. Also, except for <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="carnavalus">carnavalus</tp:taxon-name-part></tp:taxon-name></italic> Leclercq, the mid and hind legs of the females are highly modified in this group (see <xref ref-type="bibr" rid="B8">Melo and Rosa 2015</xref>). All Neotropical species also have a median protuberance on the clypeal disc, with the exception of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Trypoxylon">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="amazonicus">amazonicus</tp:taxon-name-part></tp:taxon-name></italic> Fernández &amp; Amarante, 2004. Despite lacking these features, <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species" reg="grimaldii">grimaldii</tp:taxon-name-part></tp:taxon-name></italic> sp. nov. could still turn out as sister-group of the lineage that gave rise to the Neotropical group. For example, its tear-shaped pygidial plate, with a preapical narrowing, and the broad hind femur are similar to the conditions exhibited by females of the Neotropical species. A more precise positioning of the new species must await future phylogenetic studies involving the worldwide fauna of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic>, something beyond the scope of the present contribution.</p>
      <p>The new species described here represents the seventh known apoid wasp from Dominican amber. Its discovery reinforces the bias toward crabronine inclusions in this amber. <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Tracheliodes">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> is known to nest predominantly in preexisting holes in wood (<xref ref-type="bibr" rid="B11">Zettel et al. 2004</xref>, <xref ref-type="bibr" rid="B8">Melo and Rosa 2015</xref>), a behavior that together with ant hunting make them more prone to be trapped in plant resins. Further exploration of Dominican amber accessions will certainly reveal new apoid wasps belonging to other groups besides <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Crabroninae</tp:taxon-name-part></tp:taxon-name>. Indeed, we have examined a female specimen of <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus" reg="Spilomena">Spilomena</tp:taxon-name-part></tp:taxon-name></italic> Shuckard, 1838 (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="subfamily">Pemphredoninae</tp:taxon-name-part></tp:taxon-name>) from the AMNH amber collection that remains undescribed.</p>
    </sec>
    <sec sec-type="Acknowledgements" id="SECID0EXKAC">
      <title>Acknowledgements</title>
      <p>This work started a long time ago when Caroline Chaboo brought a series of AMNH’s Dominican amber specimens to the attention of the first author. We thank her and David Grimaldi <!--PageBreak-->for the opportunity to study this material. We also would like to thank the reviewers Wojciech Pulawski and Germán Villamizar for suggestions that helped improve this article. Partial support has been provided by the Conselho Nacional de Desenvolvimento Científico e Tecnológico (CNPq) to BBR (grant 158310/2018-7) and to GARM (grant 309641/2016-0).</p>
    </sec>
  </body>
  <back>
    <ref-list>
      <title>Literature cited</title>
      <ref id="B1">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Bennett</surname><given-names>DJ</given-names></name></person-group> (<year>2010</year>) <source>Phylogenetics of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part></tp:taxon-name>, with a consideration of the evolution of predatory and nesting behaviors (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="class">Insecta</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name>). Ph.D.</source><publisher-name>thesis</publisher-name>, <publisher-loc>University of Kansas, Lawrence</publisher-loc>, <size units="page">189 pp</size>. <ext-link xlink:type="simple" ext-link-type="uri" xlink:href="https://kuscholarworks.ku.edu/handle/1808/7627">https://kuscholarworks.ku.edu/handle/1808/7627</ext-link></mixed-citation>
      </ref>
      <ref id="B2">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Bohart</surname><given-names>RR</given-names></name><name name-style="western"><surname>Menke</surname><given-names>AS</given-names></name></person-group> (<year>1976</year>) <source>Sphecid wasps of the world: A generic revision.</source><publisher-name>University of California Press</publisher-name>, <publisher-loc>Berkeley</publisher-loc>, <size units="page">695 pp</size>.</mixed-citation>
      </ref>
      <ref id="B3">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Cockerell</surname><given-names>TDA</given-names></name></person-group> (<year>1909</year>) <article-title>Descriptions of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name> from Baltic amber.</article-title><source>Schriften der Physikalisch-Ökonomischen Gesellschaft zu Königsberg</source><volume>50</volume>: <fpage>1</fpage>–<lpage>20</lpage>.</mixed-citation>
      </ref>
      <ref id="B4">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Grimaldi</surname><given-names>DA</given-names></name></person-group> (<year>1995</year>) <article-title>The age of Dominican amber.</article-title> In: <person-group><name name-style="western"><surname>Anderson</surname><given-names>KB</given-names></name><name name-style="western"><surname>Crelling</surname><given-names>JC</given-names></name></person-group> (<role>Eds</role>) <issue-title>Amber, Resinite, and Fossil Resins.</issue-title><source>American Chemical Society, Washington DC, ACS Symposium Series</source>, <fpage>203</fpage>–<lpage>217</lpage>.</mixed-citation>
      </ref>
      <ref id="B5">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Grimaldi</surname><given-names>DA</given-names></name></person-group> (<year>1996</year>) <source>Amber: window to the past. Harry N.</source><publisher-name>Abrahams</publisher-name>, <publisher-loc>New York</publisher-loc>, <size units="page">216 pp</size>.</mixed-citation>
      </ref>
      <ref id="B6">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Grimaldi</surname><given-names>DA</given-names></name><name name-style="western"><surname>Engel</surname><given-names>MS</given-names></name></person-group> (<year>2005</year>) <source>Evolution of the insects.</source><publisher-name>Cambridge University Press</publisher-name>, <publisher-loc>New York</publisher-loc>, <size units="page">722 pp</size>.</mixed-citation>
      </ref>
      <ref id="B7">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Iturralde-Vinent</surname><given-names>MA</given-names></name><name name-style="western"><surname>MacPhee</surname><given-names>RDE</given-names></name></person-group> (<year>1996</year>) <article-title>Age and paleogeographical origin of Dominican amber.</article-title><source>Science</source><volume>273</volume>: <fpage>1850</fpage>–<lpage>1852</lpage>.</mixed-citation>
      </ref>
      <ref id="B8">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Melo</surname><given-names>GAR</given-names></name><name name-style="western"><surname>Rosa</surname><given-names>BB</given-names></name></person-group> (<year>2015</year>) <article-title>A new species of the ant-hunter genus <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Tracheliodes</tp:taxon-name-part></tp:taxon-name></italic> Morawitz (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="tribe">Crabronini</tp:taxon-name-part></tp:taxon-name>) from Brazil.</article-title><source>Zootaxa</source><volume>3941</volume>: <fpage>421</fpage>–<lpage>428</lpage>. <ext-link xlink:type="simple" ext-link-type="doi" xlink:href="10.11646/zootaxa.3941.3.8">https://doi.org/10.11646/zootaxa.3941.3.8</ext-link></mixed-citation>
      </ref>
      <ref id="B9">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Penney</surname><given-names>D</given-names></name></person-group> (<year>2010</year>) <article-title>Dominican Amber.</article-title> In: <person-group><name name-style="western"><surname>Penney</surname><given-names>D</given-names></name></person-group> (<role>Ed.</role>) <issue-title>Biodiversity of fossils in amber from the major world deposits.</issue-title><source>Siri Scientific Press, Manchester</source>, <fpage>22</fpage>–<lpage>41</lpage>.</mixed-citation>
      </ref>
      <ref id="B10">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Pulawski</surname><given-names>WJ</given-names></name></person-group> (<year>2019</year>) Catalog of <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Sphecidae</tp:taxon-name-part></tp:taxon-name><italic>sensu lato.</italic><ext-link xlink:type="simple" ext-link-type="uri" xlink:href="https://www.calacademy.org/scientists/projects/catalog-of-sphecidae">https://www.calacademy.org/scientists/projects/catalog-of-sphecidae</ext-link></mixed-citation>
      </ref>
      <ref id="B11">
        <mixed-citation xlink:type="simple"><person-group><name name-style="western"><surname>Zettel</surname><given-names>H</given-names></name><name name-style="western"><surname>Ljubomirov</surname><given-names>T</given-names></name><name name-style="western"><surname>Steiner</surname><given-names>FM</given-names></name><name name-style="western"><surname>Schlick-Steiner</surname><given-names>BC</given-names></name><name name-style="western"><surname>Grabenweger</surname><given-names>G</given-names></name><name name-style="western"><surname>Wiesbauer</surname><given-names>H</given-names></name></person-group> (<year>2004</year>) <article-title>The European ant hunters <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">Tracheliodes</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">curvitarsus</tp:taxon-name-part></tp:taxon-name></italic> and <italic><tp:taxon-name><tp:taxon-name-part taxon-name-part-type="genus">T.</tp:taxon-name-part> <tp:taxon-name-part taxon-name-part-type="species">varus</tp:taxon-name-part></tp:taxon-name></italic> (<tp:taxon-name><tp:taxon-name-part taxon-name-part-type="order">Hymenoptera</tp:taxon-name-part></tp:taxon-name>: <tp:taxon-name><tp:taxon-name-part taxon-name-part-type="family">Crabronidae</tp:taxon-name-part></tp:taxon-name>): taxonomy, species discrimination, distribution, and biology.</article-title><source>Myrmecologische Nachrichten</source><volume>6</volume>: <fpage>39</fpage>–<lpage>47</lpage>.</mixed-citation>
      </ref>
    </ref-list>
  </back>
</article>
