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    <front>
        <journal-meta>
            <journal-id journal-id-type="publisher-id">ZooKeys</journal-id>
            <journal-title-group>
                <journal-title xml:lang="en">ZooKeys</journal-title>
                <abbrev-journal-title xml:lang="en">ZooKeys</abbrev-journal-title>
            </journal-title-group>
            <issn pub-type="ppub">1313-2989</issn>
            <issn pub-type="epub">1313-2970</issn>
            <publisher>
                <publisher-name>Pensoft Publishers</publisher-name>
            </publisher>
        </journal-meta>
        <article-meta>
            <article-id pub-id-type="doi">10.3897/zookeys.369.6561</article-id>
            <title-group>
                <article-title>Two new species of <italic>Archaeohelorus</italic> (Hymenoptera, Proctotrupoidea, Heloridae) from the Middle Jurassic of China</article-title>
            </title-group>
            <contrib-group>
                <contrib contrib-type="author" xlink:type="simple">
                    <name name-style="western">
                        <surname>Shi</surname>
                        <given-names>Xiaoqing</given-names>
                    </name>
                    <xref ref-type="aff" rid="A1">1</xref>
                    <uri content-type="zoobank" xlink:type="simple">http://zoobank.org/A799217E-63CF-4EE8-BB7D-3BB4D8A65E0D</uri>
                </contrib>
                <contrib contrib-type="author" xlink:type="simple">
                    <name name-style="western">
                        <surname>Zhao</surname>
                        <given-names>Yunyun</given-names>
                    </name>
                    <xref ref-type="aff" rid="A1">1</xref>
                    <uri content-type="zoobank" xlink:type="simple">http://zoobank.org/C1AE36DE-8D4C-4DA5-BD96-E435A584A5D3</uri>
                </contrib>
                <contrib contrib-type="author" xlink:type="simple">
                    <name name-style="western">
                        <surname>Shih</surname>
                        <given-names>Chungkun</given-names>
                    </name>
                    <xref ref-type="aff" rid="A1">1</xref>
                    <uri content-type="zoobank" xlink:type="simple">http://zoobank.org/6FC8E402-B9F2-48B5-B3F2-71B91C09CA7C</uri>
                </contrib>
                <contrib contrib-type="author" xlink:type="simple">
                    <name name-style="western">
                        <surname>Ren</surname>
                        <given-names>Dong</given-names>
                    </name>
                    <xref ref-type="aff" rid="A1">1</xref>
                    <uri content-type="zoobank" xlink:type="simple">http://zoobank.org/D507ABBD-6BA6-43C8-A1D5-377409BD3049</uri>
                </contrib>
            </contrib-group>
            <aff id="A1">
                <label>1</label>College of Life Sciences, Capital Normal University, 105 Xisanhuanbeilu, Haidian District, Beijing 100048, China
            </aff>
            <author-notes>
                <fn fn-type="corresp">
                    <p>Corresponding author: Dong Ren (<email xlink:type="simple">rendong@mail.cnu.edu.cn</email>)</p>
                </fn>
                <fn fn-type="edited-by">
                    <p>Academic editor: Jes Rust</p>
                </fn>
            </author-notes>
            <pub-date pub-type="collection">
                <year>2014</year>
            </pub-date>
            <pub-date pub-type="epub">
                <day>13</day>
                <month>1</month>
                <year>2014</year>
            </pub-date>
            <issue>369</issue>
            <fpage>49</fpage>
            <lpage>59</lpage>
            <history>
                <date date-type="received">
                    <day>4</day>
                    <month>11</month>
                    <year>2013</year>
                </date>
                <date date-type="accepted">
                    <day>24</day>
                    <month>12</month>
                    <year>2013</year>
                </date>
            </history>
            <permissions>
                <copyright-statement>Xiaoqing Shi, Yunyun Zhao, Chungkun Shih, Dong Ren</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/57D43135-2C67-46A4-BB3A-0A163B5A025D</self-uri>
            <abstract>
                <label>Abstract</label>
                <p>Two new fossil species, <italic>Archaeohelorus polyneurus</italic> <bold>sp. n.</bold> and <italic>A. tensus</italic> <bold>sp. n.</bold>, assigned to the genus <italic>Archaeohelorus</italic> Shih, Feng &amp; Ren, 2011 of Heloridae (Hymenoptera), are reported from the late Middle Jurassic, Jiulongshan Formation of Inner Mongolia, China. Based on the well-preserved forewings and hind wings of these specimens, the diagnosis of the <italic>Archaeohelorus</italic> is emended: forewing 2cu-a intersecting Cu and Rs+M at the same point or postfurcal, and hind wing may have tubular veins C, Sc+R, R, Rs, M+Cu, M and Cu distinct, or simplified venation. The new findings also elucidate the evolutionary trend of forewing and hind wing venation and body size for the Heloridae from the late Middle Jurassic to now.</p>
            </abstract>
            <kwd-group>
                <label>Keywords</label>
                <kwd>Fossil wasps</kwd>
                <kwd>Heloridae</kwd>
                <kwd><italic>Archaeohelorus</italic></kwd>
                <kwd>Middle Jurassic</kwd>
                <kwd>China</kwd>
                <!--PageBreak-->
            </kwd-group>
        </article-meta>
    </front>
    <body>
        <sec sec-type="Introduction">
            <title>Introduction</title>
            <p><tp:taxon-name>Proctotrupoidea</tp:taxon-name> Latreille, 1802, including 11 extant families, is a significant group within <tp:taxon-name>Hymenoptera</tp:taxon-name> for their long evolutionary history, special morphology and diversity (<xref ref-type="bibr" rid="B2">Grimaldi and Engel 2005</xref>). Most extant species of <tp:taxon-name>Proctotrupoidea</tp:taxon-name> are small wasps except for the giants of the family <tp:taxon-name>Pelecinidae</tp:taxon-name> (<xref ref-type="bibr" rid="B20">Shih et al. 2010</xref>). <tp:taxon-name>Heloridae</tp:taxon-name>, a small family in <tp:taxon-name>Proctotrupoidea</tp:taxon-name>, have the earliest fossil records from the late Middle Jurassic (<xref ref-type="bibr" rid="B21">Shih et al. 2011</xref>).</p>
            <p>Up to date, fossil <tp:taxon-name>Heloridae</tp:taxon-name> contains 8 genera and 12 species, which have been summarized by <xref ref-type="bibr" rid="B19">Shi et al. (2012)</xref>. These species have been described from the late Middle Jurassic of Daohugou Ningcheng, China (<xref ref-type="bibr" rid="B21">Shih et al. 2011</xref>); the Late Jurassic of Karatau, Russia (<xref ref-type="bibr" rid="B17">Rohdendorf 1938</xref>), and of Laiyang, China (<xref ref-type="bibr" rid="B30">Zhang 1992</xref>); the Early Cretaceous of Turga, Russia (<xref ref-type="bibr" rid="B12">Rasnitsyn 1990</xref>), of Beipiao, China (<xref ref-type="bibr" rid="B29">Zhang and Zhang 2001</xref>; <xref ref-type="bibr" rid="B19">Shi et al. 2012</xref>); and of Gurvan-Ereny-Nuru, Western Mongolia (<xref ref-type="bibr" rid="B11">Rasnitsyn 1986</xref>).</p>
            <p>So far, 7 species of <tp:taxon-name>Heloridae</tp:taxon-name> from China have been reported, including <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011; <italic><tp:taxon-name>Gurvanhelorus beipiaoensis</tp:taxon-name></italic> Shi, Shih &amp; Ren, 2012; <italic><tp:taxon-name>Protocyrtus validus</tp:taxon-name></italic> Zhang &amp; Zhang, 2001; <italic><tp:taxon-name>Sinohelorus elegans</tp:taxon-name></italic> Shi, Shih &amp; Ren, 2012; <italic><tp:taxon-name>Spherogaster coronata</tp:taxon-name></italic> Zhang &amp; Zhang, 2001; and <italic><tp:taxon-name>Spherogaster saltatrix</tp:taxon-name></italic> Shi, Shih &amp; Ren, 2012.</p>
            <p>Extant helorids contains only one genus, <italic><tp:taxon-name>Helorus</tp:taxon-name></italic> Latreille, 1802, with 12 known valid species mostly in the Holarctic Region. They are parasitoids of larvae of chrysopid lacewings (<tp:taxon-name>Neuroptera</tp:taxon-name>: <tp:taxon-name>Chrysopidae</tp:taxon-name>: <tp:taxon-name>Chrysopinae</tp:taxon-name>: <italic><tp:taxon-name>Chrysopa</tp:taxon-name></italic> species) (<xref ref-type="bibr" rid="B23">van Achterberg 2006</xref>). Among the known species, only <italic><tp:taxon-name>Helorus chinensis</tp:taxon-name></italic> He, 1992 was described from China (<xref ref-type="bibr" rid="B4">He 1992</xref>).</p>
            <p>Recently, we collected two well-preserved fossil specimens referable to <tp:taxon-name>Heloridae</tp:taxon-name> from the late Middle Jurassic Jiulongshan Formation at Daohugou Village, Ningcheng County, Inner Mongolia, China. Based on these new findings, the diagnosis of <italic><tp:taxon-name>Archaeohelorus</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011 is emended and two new species, <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n. and <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> sp. n., are described. This is the second report of <tp:taxon-name>Heloridae</tp:taxon-name> in the late Middle Jurassic.</p>
            <p>The age of the Daohugou fossil-bearing beds is interpreted to be late Middle Jurassic (<italic>ca</italic> 165 Ma; <xref ref-type="bibr" rid="B16">Ren et al. 2010</xref>). This deposit is interpreted to have accumulated in streams and lakes within a humid and warm-temperate climate (<xref ref-type="bibr" rid="B14">Ren et al. 2002</xref>). It is rich in well-preserved fossils, especially a high level of insect diversity have been reported including <tp:taxon-name>Ephemeroptera</tp:taxon-name> (<xref ref-type="bibr" rid="B6">Huang et al. 2008</xref>), <tp:taxon-name>Odonata</tp:taxon-name> (<xref ref-type="bibr" rid="B7">Li et al. 2011</xref>), <tp:taxon-name>Plecoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B9">Liu et al. 2011</xref>), <tp:taxon-name>Blattodea</tp:taxon-name> (<xref ref-type="bibr" rid="B28">Wei et al. 2012</xref>), <tp:taxon-name>Orthoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B3">Gu et al. 2012</xref>), <tp:taxon-name>Homoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B24">Wang et al. 2012</xref>), <tp:taxon-name>Heteroptera</tp:taxon-name> (<xref ref-type="bibr" rid="B10">Lu et al. 2011</xref>), <tp:taxon-name>Neuroptera</tp:taxon-name> (<xref ref-type="bibr" rid="B26">Wang et al. 2010</xref>; <xref ref-type="bibr" rid="B18">Shi et al. 2011</xref>), <tp:taxon-name>Raphidioptera</tp:taxon-name> (Engel &amp; Ren, 2008), <tp:taxon-name>Coleoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B22">Tan et al. 2012</xref>), <tp:taxon-name>Mecoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B15">Ren et al. 2009</xref>; <xref ref-type="bibr" rid="B24">Wang et al. 2012</xref>), <tp:taxon-name>Hymenoptera</tp:taxon-name> (<xref ref-type="bibr" rid="B24">Wang et al. 2012</xref>), and <tp:taxon-name>Diptera</tp:taxon-name> (<xref ref-type="bibr" rid="B8">Liu et al. 2012</xref>).</p>
        </sec>
        <sec sec-type="materials|methods">
            <title>Materials and methods</title>
            <p>All the materials have been collected near Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China; the late Middle Jurassic (Bathonian-Call<!--PageBreak-->ovian boundary, 165 Ma). All fossil specimens are housed in the Key Lab of Insect Evolution &amp; Environmental Changes, College of Life Sciences, Capital Normal University, Beijing, China.</p>
            <p>The specimens were examined dry or under alcohol using a M165 C dissecting microscope (Leica) and are illustrated with the aid of a drawing tube attachment. The figures were drawn by Adobe Photoshop CS5 and CorelDraw 12.0. Morphological terminology and the system used here follow those of <xref ref-type="bibr" rid="B5">Huber and Sharkey (1993)</xref> and <xref ref-type="bibr" rid="B13">Rasnitsyn and Zhang (2010)</xref>.</p>
        </sec>
        <sec sec-type="Systematic Paleontology">
            <title>Systematic Paleontology</title>
            <sec sec-type="Class Insecta L., 1758">
                <title>Class <tp:taxon-name>Insecta</tp:taxon-name> L., 1758<break/>
                Order <tp:taxon-name>Hymenoptera</tp:taxon-name> L., 1758<break/>
                Suborder <tp:taxon-name>Apocrita</tp:taxon-name> Gerstaecker, 1867<break/>
                Superfamily <tp:taxon-name>Proctotrupoidea</tp:taxon-name> Latreille, 1802<break/>
                Family <tp:taxon-name>Heloridae</tp:taxon-name> Foerster, 1856<break/>
                Subfamily <tp:taxon-name>Mesohelorinae</tp:taxon-name> Rasnitsyn, 1990</title>
                <tp:taxon-treatment>
                    <tp:nomenclature>
                        <label>Genus</label>
                        <tp:taxon-name>
                            <tp:taxon-name-part taxon-name-part-type="genus">Archaeohelorus</tp:taxon-name-part>
                            <object-id xlink:type="simple">http://species-id.net/wiki/Archaeohelorus</object-id>
                        </tp:taxon-name>
                        <tp:taxon-authority>Shih, Feng &amp; Ren, 2011</tp:taxon-authority>
                    </tp:nomenclature>
                    <tp:treatment-sec sec-type="Type species">
                        <title>Type species.</title>
                        <p><italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Emended diagnosis">
                        <title>Emended diagnosis.</title>
                        <p>Forewing 2cu-a intersecting Cu and Rs+M at the same point or postfurcal. Hind wing may have tubular veins C and Sc+R separated at base, R developed, M+Cu robust and forking at the basal part, M and Cu distinct, or simplified venation.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Species included">
                        <title>Species included.</title>
                        <p><italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011, <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n. and <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> sp. n.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Remarks">
                        <title>Remarks.</title>
                        <p>This genus was established by <xref ref-type="bibr" rid="B21">Shih et al. 2011</xref> based on a holotype, allotype and six paratypes from the Middle Jurassic of Daohugou, Inner Mongolia, China. Due to lack of discernible hind wings on the fossils of holotype and allotype, the hind wing venation was not described. With the new forewing and hind wing venational information on our new materials, we emended the generic diagnosis.</p>
                    </tp:treatment-sec>
                </tp:taxon-treatment>
                <tp:taxon-treatment>
                    <tp:nomenclature>
                        <tp:taxon-name>
                            <tp:taxon-name-part taxon-name-part-type="genus">Archaeohelorus</tp:taxon-name-part>
                            <tp:taxon-name-part taxon-name-part-type="species">polyneurus</tp:taxon-name-part>
                            <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/BFECD83C-516A-4F08-BBB3-72A5EB6DFF41</object-id>
                            <object-id xlink:type="simple">http://species-id.net/wiki/Archaeohelorus_polyneurus</object-id>
                        </tp:taxon-name>
                        <tp:taxon-status xlink:type="simple">sp. n.</tp:taxon-status>
                        <xref ref-type="fig" rid="F1">Figures 1</xref>
                        <xref ref-type="fig" rid="F2">, 2</xref>
                    </tp:nomenclature>
                    <tp:treatment-sec sec-type="Etymology">
                        <title>Etymology.</title>
                        <p>The specific nameis from Greek word “<italic>polyneurus</italic>”, means “many veins”, referring to the complete venation of the hind wing preserved.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Type material">
                        <title>Type material.</title>
                        <p>Holotype, CNU-HYM-NN2012052, dorsal view, gender unknown. A well-preserved body with almost complete forewings and hind wings and <!--PageBreak-->part of legs, but head missing. Paratype: CNU-HY-NN2008010, dorsal view, a well-preserved almost complete body with forewings and right hind wing and part of legs, previously mis-identified as a paratype of <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Locality and age">
                        <title>Locality and age.</title>
                        <p>Jiulongshan Formation, Middle Jurassic, Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Diagnosis">
                        <title>Diagnosis.</title>
                        <p>In forewing, 2cu-a intersecting Cu and Rs+M at the same point and cell r obtuse-angled triangular (vs. approximately right-angled triangular in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>). Hind wing with Sc+R confluent with C and extended to costal margin, R developed. M+Cu distinct and forking at the basal part, M and Cu robust.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Remarks">
                        <title>Remarks.</title>
                        <p>Upon further examination, we found that a paratype of <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>, CNU-HYM-NN2008010, has forewing 2cu-a intersecting Cu and Rs+M at the same point and cell r obtuse-angled triangular (vs. approximately right-angled triangular in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>); and hind wing with tubular vein C parallel with Sc+R at base, M+Cu robust and forking at the basal section, M long, 1-Cu, 2-Cu distinct. These venational characters are consistent with the diagnostic characters of <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n., hence we transfer CNU-HYM-NN2008010 as a paratype of <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Description of holotype">
                        <title>Description of holotype.</title>
                        <p>A medium-sized body with both forewings and hind wings well-preserved, but, without head (<xref ref-type="fig" rid="F1">Fig. 1</xref>). Forewing broad. Mesosoma suboval, nearly 1.35 times as long as wide; mesoscutum trapezoidal with notauli distinct and <!--PageBreak-->concave; tegula triangular; scutellum broad with two rows of pits; metanotum relatively wide with plenty of pits; propodeum transverse, 2.89 times as broad as long.</p>
                        <fig id="F1" position="float" orientation="portrait">
                            <label>Figure 1.</label>
                            <caption><p>Holotype of <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n. CNU-HYM-NN-2012052. <bold>A</bold> Photo <bold>B</bold> line drawing <bold>C</bold> line drawing of forewing <bold>D</bold> line drawing of hind wing. Scale bars: 1 mm. (Online figure in color.)</p></caption>
                            <graphic xlink:href="ZooKeys-369-049-g001.jpg" position="float" orientation="portrait" xlink:type="simple"></graphic>
                        </fig>
                        <p>Metasoma suboval with six segments; first metasomal segment transverse, second segment with several longitudinal ridges; third and fourth segments trapezoidal with shorter longitudinal ridges anteriorly; other ones smooth; the end of terminal segment not preserved.</p>
                        <p>Forewing broad and subtriangular. Pterostigma long and acute apically, not widened beyond mid-length. C robust and extending to apex of forewing. 2r-rs arising from basal one third of pterostigma, slightly oblique apicad, slightly longer than pterostigmal width. R robust, cell r obtuse-angled triangular and closed by R, M+Cu, 1-Rs and 1-M; Rs straight and intersecting with distal part of C. Cell 1+2r six-sided and surrounded by R, 1-Rs, 1-Rs+M, 2-Rs+M, 2-Rs, and 2r-rs. M+Cu straight and distinct; M and Cu distinct, M straight, Cu break after 1m-cu. 1-Rs as long as 1-M and slightly inclined toward wing base. Cell 1mcu small and subtriangular. M and Rs branching at 30% from 1m-cu of the length between 1m-cu and 2r-rs. 1cu-a and 2cu-a distinct and reaching A. 1cu-a in line with 1-M; and 2cu-a intersecting Cu and Rs+M at the same point.</p>
                        <p>Hind wing with tubular veins C parallel with Sc+R at base; R developed and Rs preserved; M+Cu robust and forking at the basal section, M and 1-Cu long and distinct, 2-Cu long.</p>
                        <p>Right foreleg with only coxa and femur partially preserved. Right midleg with partial trochanter, spindle-shaped femur and partial tibia preserved. Right hindleg with partial coxa, trochanter trapezoidal and small, robust spindle-shaped femur, long tibia <!--PageBreak-->swollen distally with spurs, tarsi with five segments, basitarsus longest and two claws fixing the end of pretarsus.</p>
                        <p><bold>Measurements (in mm).</bold> Mesosoma length 3.36, width 2.48; metasoma &gt; 3.38 long; lengths of the first to fifth metasomal segments are 0.34, 0.51, 0.72, 0.94, and 0.55; forewing length 6.52, width &gt; 3.05.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Description of paratype">
                        <title>Description of paratype.</title>
                        <p>Body medium-sized (<xref ref-type="fig" rid="F2">Fig. 2</xref>). Head oval and antennae filiform and thick, scape swollen and bell-shaped, only first and second flagellomeres preserved. Mesosoma, forewing and metasoma same as holotype, the end of terminal segment not preserved. Hind wing with tubular vein C parallel with Sc+R at base, M+Cu robust and forking at the basal section, M long, 1-Cu, 2-Cu distinct. Left foreleg and right midleg with only coxa and femur partially preserved. Left midleg with femur robust and tibia long, left hindleg with femur robust and spindle-shaped, long tibia swollen distally. Right hindleg with femur robust and spindle-shaped, tibia partially preserved.</p>
                        <fig id="F2" position="float" orientation="portrait">
                            <label>Figure 2.</label>
                            <caption><p>Paratype of <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n. CNU-HYM-NN-2008010. <bold>A</bold> Photo <bold>B</bold> line drawing <bold>C</bold> line drawing of forewing <bold>D</bold> lLine drawing of hind wing. Scale bars: 1 mm. (Online figure in color.)</p></caption>
                            <graphic xlink:href="ZooKeys-369-049-g002.jpg" position="float" orientation="portrait" xlink:type="simple"></graphic>
                        </fig>
                        <p><bold>Measurements (in mm).</bold> Body length &gt; 14.6; head length 1.89, width 1.53; mesosoma length 3.58, width 2.52; metasoma length 3.68; forewing length &gt; 6.54, width &gt; 2.88.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Comparison">
                        <title>Comparison.</title>
                        <p>This species can be assigned to <italic><tp:taxon-name>Archaeohelorus</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011 by its metasoma with six segments and the first segment narrow and transverse. Forewing 1-Rs as long as 1-M. Cell 1mcu small and subtriangular, 1cu-a in line with 1-M; and 2cu-a intersecting Cu and Rs+M at the same point. Compared with <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>, <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> has preserved hind wing with Sc+R confluent with C and extended to costal margin, R developed. M+Cu distinct and forking at the basal part, M and Cu robust; in forewing, cell r obtuse-angled triangular (vs. approximately right-angled triangular in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>). Besides, <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> has a much larger body size than <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>, length of forewing 6.52 mm or &gt; 6.54 mm (vs. 3.17 mm in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>).</p>
                    </tp:treatment-sec>
                </tp:taxon-treatment>
                <tp:taxon-treatment>
                    <tp:nomenclature>
                        <tp:taxon-name>
                            <tp:taxon-name-part taxon-name-part-type="genus">Archaeohelorus</tp:taxon-name-part>
                            <tp:taxon-name-part taxon-name-part-type="species">tensus</tp:taxon-name-part>
                            <object-id content-type="zoobank" xlink:type="simple">http://zoobank.org/84E4B637-E6E6-4799-929E-1536DA5213A5</object-id>
                            <object-id xlink:type="simple">http://species-id.net/wiki/Archaeohelorus_tensus</object-id>
                        </tp:taxon-name>
                        <tp:taxon-status xlink:type="simple">sp. n.</tp:taxon-status>
                        <xref ref-type="fig" rid="F3">Figure 3</xref>
                    </tp:nomenclature>
                    <tp:treatment-sec sec-type="Etymology">
                        <title>Etymology.</title>
                        <p>The specific name “<italic>tensus</italic>” means stretching and long, referring to the shape of the forewing.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Type material">
                        <title>Type material.</title>
                        <p>Holotype, CNU-HYM-NN2012056p/c, part and counterpart, dorsal view, male. A well-preserved almost complete body with antenna, forewings and part of legs and partial hind wings.</p>
                        <fig id="F3" position="float" orientation="portrait">
                            <label>Figure 3.</label>
                            <caption><p>Holotype of <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> sp. n. CNU-HYM-NN-2012056 p/c, part and counterpart. <bold>A</bold> Photo of part <bold>B</bold> photo of counterpart <bold>C</bold> line drawing of part <bold>D</bold> line drawing of forewing <bold>E</bold> terminal segments of counterpart (under alcohol). Scale bars: 1 mm. (Online figure in color.)</p></caption>
                            <graphic xlink:href="ZooKeys-369-049-g003.jpg" position="float" orientation="portrait" xlink:type="simple"></graphic>
                        </fig>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Locality and age">
                        <title>Locality and age.</title>
                        <p>Jiulongshan Formation, Middle Jurassic, Daohugou Village, Shantou Township, Ningcheng County, Inner Mongolia, China.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Diagnosis">
                        <title>Diagnosis.</title>
                        <p>Forewing 2cu-a postfurcal with intersection of Cu and Rs+M. First abscissa of Rs (1-Rs) and basal section of M (1-M) arched toward basal of wing. Hind wing with tubular veins C, Sc+R, and Rs preserved.</p>
                        <!--PageBreak-->
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Description">
                        <title>Description.</title>
                        <p>A smalladult specimen with a total body length of 4.4 mm. Head suboval and large relative to mesosoma in width. Eyes large, and located at the both sides of the head.</p>
                        <p>Antennae filiform, thick, with 6 segments preserved on left and 5 segments preserved on right. Scape swollen and bell-shaped; pedicel short and quadrate; subsequent flagellomeres not well preserved.</p>
                        <p>Mesosoma subhexagonal and broader than head; pronotum narrower than head, short, probably covered by mesonotum; mesoscutum distorted in the middle and trapezoidal with notauli not distinct but preserved; tegula large and subtriangular; scutellum round with pits anteriorly; propodeum narrow, transverse and areolated.</p>
                        <p>Metasoma suboval with seven segments; first metasomal segment transverse with several longitudinal ridges; second segment trapezoidal with less longitudinal ridges anteriorly; other ones smooth; the male terminalia triangular, partially covered by the previous segment.</p>
                        <p>Forewing broad and subtriangular. Pterostigma long and acute apically, not widened beyond mid-length. C robust and extending near the apex of forewing. 2r-rs arising from basal one third of pterostigma, slightly longer than the width of pterostigma, and slightly oblique apicad. R robust and cell r closed with C, R and 1-RS. Rs straight and intersecting with distal part of C. Cell 1+2r longer and narrower relatively and six-sided <!--PageBreak-->surrounded by R, 1-Rs, 1-Rs+M, 2-Rs+M, 2-Rs, and 2r-rs. M+Cu straight and distinct; M and Cu distinct, almost straight. 1-Rs as long as 1-M and slightly inclined toward wing base. Cell 1mcu small, subtriangular and relatively slender with 2-M+Cu 4.3 times as long as 1-M. M and Rs branching at 30% from 1m-cu of the length between 1m-cu and 2r-rs. 1cu-a and 2cu-a distinct and reaching A. 1cu-a in line with 1-M; and 2cu-a postfurcal with Rs+M distinctly.</p>
                        <p>Hind wing, with tubular veins C parallel with Sc+R at base, Rs short, intersecting distal part of C.</p>
                        <p>Left foreleg, midleg and hindleg and right foreleg with only coxa and femur partially preserved, hind femur much thicker, nearly three times as long as wide; right midleg with partial spindle-shaped femur and partial tibia preserved, right hindleg with partial coxa, trapezoidal trochanter, robust spindle-shaped femur and relatively thin tibia and some parts of tarsus preserved.</p>
                        <p><bold>Measurements (in mm):</bold> Body length 4.67, head length1.39, width 0.68, mesosoma length 1.68, width 1.36; metasoma 2.23 long; lengths of metasomal segments are 0.20, 0.39, 0.48, 0.38, 0.25 and 0.32; forewing length &gt; 2.96, width &gt; 1.36.</p>
                    </tp:treatment-sec>
                    <tp:treatment-sec sec-type="Remarks and comparison">
                        <title>Remarks and comparison.</title>
                        <p>This species is assigned to <italic><tp:taxon-name>Archaeohelorus</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011 by its seven separated metasomal segments, the first metasomal segment transverse with several longitudinal ridges. Forewing 1-Rs as long as 1-M. Cell 1mcu small and subtriangular, 1cu-a in line with 1-M. Compared with <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>, <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> has preserved hind wing with tubular veins C, Sc+R, R, and Rs preserved; in forewing, 2cu-a distinctly postfurcal with intersection of Cu and Rs+M (vs. 2cu-a intersecting Cu and Rs+M at the same point in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>); cell 1mcu distinctly slender, 4.3 times as long as wide (vs. 2.4 times in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>); 1-Rs and 1-M arched toward basal of wing (vs. 1-Rs and 1-M straight in <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic>). It also differs from <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> by its postfurcal 2cu-a, slender cell 1mcu.</p>
                    </tp:treatment-sec>
                </tp:taxon-treatment>
            </sec>
        </sec>
        <sec sec-type="Discussion">
            <title>Discussion</title>
            <p>In the vast and extensive Daohugou fossil insect collection (&gt;200,000 insect fossil specimens) at the Capital Normal University, only ten helorids are collected so far: two are <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> sp. n., one is <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> sp. n., and seven specimens are previously described <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic> Shih, Feng &amp; Ren, 2011. It is likely that helorids might have been a very small group in the Middle Jurassic.</p>
            <p>As a relict family, helorids have survived from the Middle Jurassic to now. Among all the fossil specimens, only <italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> has distinct and complicated hind wing venation, which were unknown before. Therefore, the new findings are important supplement to helorid record and suggest that the trend for the morphological evolution of the <tp:taxon-name>Heloridae</tp:taxon-name> is as follows:</p>
            <p>In the Middle Jurassic, <!--PageBreak--><italic><tp:taxon-name>Archaeohelorus polyneurus</tp:taxon-name></italic> had forewing cell r obtuse-angled triangle, and hind wing venation clear and more complex (C and Sc+R separated at base, M+Cu short and robust, forking at the basal part, M and Cu distinct) and a much larger body (forewing length &gt; 6.52mm). On the other hand, <italic><tp:taxon-name>Archaeohelorus hoi</tp:taxon-name></italic> and <italic><tp:taxon-name>Archaeohelorus tensus</tp:taxon-name></italic> had forewing cell r approximately right-angled triangular, and hind with simplified venation (only C and Sc+R and/or R preserved) and smaller body (forewing length ≈ 3 mm). In the Late Jurassic to the Early Cretaceous, <italic><tp:taxon-name>Spherogaster coronata</tp:taxon-name></italic> Zhang &amp; Zhang, 2001 had hind wing with only C, R and M present, costal area extremely narrow, and a much larger body size with forewing length of 12 mm. In extant helorids, forewing cell r approximately right-angled triangular, and hind wings have tubular veins C and Sc+R and nebulosus veins of M+Cu, M, Cu and A (Goulet and Huber 1993). The body size is small with forewing about 3.3 mm long (<xref ref-type="bibr" rid="B23">van Achterberg 2006</xref>).</p>
        </sec>
    </body>
    <back>
        <ack>
            <title>Acknowledgments</title>
            <p>We are sincerely grateful to Qiang Yang and Taiping Gao (College of Life Sciences, Capital Normal University) for their valuable comments and suggestion on the manuscript. This work was supported by the National Basic Research Program of China 973 Program Grant 2012CB821906; National Natural Science Foundation of China Grants, 31230065 and 41272006; Great Wall Scholar and KEY project of Beijing Municipal Commission of Education (grant KZ201310028033), Program for Changjiang Scholars and Innovative Research Team in University.</p>
        </ack>
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