<?xml version="1.0" encoding="UTF-8"?><article article-type="normal" xml:lang="en">
   <front>
      <journal-meta>
         <journal-id journal-id-type="publisher-id">PALEVO</journal-id>
         <issn>1631-0683</issn>
         <publisher>
            <publisher-name>Elsevier</publisher-name>
         </publisher>
      </journal-meta>
      <article-meta>
         <article-id pub-id-type="pii">S1631-0683(16)30091-4</article-id>
         <article-id pub-id-type="doi">10.1016/j.crpv.2016.08.005</article-id>
         <article-categories>
            <subj-group subj-group-type="type">
               <subject>Research article</subject>
            </subj-group>
            <subj-group subj-group-type="heading">
               <subject>General Palaeontology, Systematics and Evolution (Invertebrate Palaeontology)</subject>
            </subj-group>
            <series-title>Invertebrate palaeontology</series-title>
         </article-categories>
         <title-group>
            <article-title>
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., a new damselfly (Odonata: Zygoptera: Platycnemididae) from mid-Cretaceous Burmese amber</article-title>
            <trans-title-group xml:lang="fr">
               <trans-title>
                  <italic>Palaeodisparoneura cretacica</italic> sp. nov., une nouvelle demoiselle (Odonata : Zygoptera : Platycnemididae) de l’ambre birman du Cretacé moyen</trans-title>
            </trans-title-group>
         </title-group>
         <contrib-group content-type="authors">
            <contrib contrib-type="author" corresp="yes">
               <name>
                  <surname>Zheng</surname>
                  <given-names>Daran</given-names>
               </name>
               <email>dranzheng@gmail.com</email>
               <xref rid="aff0005" ref-type="aff">
                  <sup>a</sup>
               </xref>
               <xref rid="aff0010" ref-type="aff">
                  <sup>b</sup>
               </xref>
            </contrib>
            <contrib contrib-type="author">
               <name>
                  <surname>Wang</surname>
                  <given-names>Bo</given-names>
               </name>
               <xref rid="aff0005" ref-type="aff">
                  <sup>a</sup>
               </xref>
               <xref rid="aff0015" ref-type="aff">
                  <sup>c</sup>
               </xref>
            </contrib>
            <contrib contrib-type="author" corresp="yes">
               <name>
                  <surname>Chang</surname>
                  <given-names>Su-Chin</given-names>
               </name>
               <email>suchin@hku.hk</email>
               <xref rid="aff0010" ref-type="aff">
                  <sup>b</sup>
               </xref>
            </contrib>
            <aff-alternatives id="aff0005">
               <aff>
                  <label>a</label> State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, 39 East Beijing Road, 210008 Nanjing, China</aff>
               <aff>
                  <label>a</label>
                  <institution>State Key Laboratory of Palaeobiology and Stratigraphy, Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences</institution>
                  <addr-line>39 East Beijing Road</addr-line>
                  <city>Nanjing</city>
                  <postal-code>210008</postal-code>
                  <country>China</country>
               </aff>
            </aff-alternatives>
            <aff-alternatives id="aff0010">
               <aff>
                  <label>b</label> Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region, China</aff>
               <aff>
                  <label>b</label>
                  <institution>Department of Earth Sciences, The University of Hong Kong, Hong Kong Special Administrative Region</institution>
                  <country>China</country>
               </aff>
            </aff-alternatives>
            <aff-alternatives id="aff0015">
               <aff>
                  <label>c</label> Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences, 1, Beichen West Road, 100101 Beijing, China</aff>
               <aff>
                  <label>c</label>
                  <institution>Key Laboratory of Zoological Systematics and Evolution, Institute of Zoology, Chinese Academy of Sciences</institution>
                  <addr-line>1, Beichen West Road</addr-line>
                  <city>Beijing</city>
                  <postal-code>100101</postal-code>
                  <country>China</country>
               </aff>
            </aff-alternatives>
         </contrib-group>
         <pub-date-not-available/>
         <volume>16</volume>
         <issue>3</issue>
         <issue-id pub-id-type="pii">S1631-0683(17)X0003-1</issue-id>
         <fpage seq="0" content-type="normal">235</fpage>
         <lpage content-type="normal">240</lpage>
         <history>
            <date date-type="received" iso-8601-date="2016-03-27"/>
            <date date-type="accepted" iso-8601-date="2016-08-15"/>
         </history>
         <permissions>
            <copyright-statement>© 2016 Académie des sciences. Published by Elsevier B.V. All rights reserved.</copyright-statement>
            <copyright-year>2016</copyright-year>
            <copyright-holder>Académie des sciences</copyright-holder>
         </permissions>
         <self-uri xmlns:xlink="http://www.w3.org/1999/xlink" content-type="application/pdf" xlink:href="main.pdf">
                        Full (PDF)
                    </self-uri>
         <abstract abstract-type="author">
            <p id="spar0005">Abundant odonatans have been discovered from mid-Cretaceous Burmese amber, and Burma has played an important role in early damselfly diversification during the mid-Cretaceous. In this paper, a new damselfly, <italic>Palaeodisparoneura cretacica</italic> sp. nov., is described from Burmese amber. It is the second species of the extinct genus <italic>Palaeodisparoneura</italic> Poinar, Bechly et Buckley, 2010. <italic>P. cretacica</italic> sp. nov. differs from <italic>P. burmanica</italic> Poinar, Bechly et Buckley, 2010 in having more postnodal and postsubnodal crossveins, the base of IR1 being more cells distal of the base of RP2, a hyaline pterostigma and a longer RP3/4. Our find increases the diversity of damselflies during the mid-Cretaceous.</p>
         </abstract>
         <trans-abstract abstract-type="author" xml:lang="fr">
            <p id="spar0010">D’abondants Odonates ont été découverts dans l’ambre birman du Crétacé moyen, et la Birmanie a joué un rôle important dans la diversification précoce des demoiselles pendant le Crétacé moyen. Dans cet article, une nouvelle demoiselle, <italic>Palaeodisparoneura</italic> <italic>cretacia</italic> sp. nov., est décrite dans l’ambre birman. C’est la seconde espèce du genre éteint <italic>Palaeodisparoneura</italic> Poinar, Bechly et Buckley, 2010. <italic>P. cretacia</italic> sp. nov. diffère de <italic>P. burmanica</italic> Poinar, Bechly et Buckley, 2010 par l’existence d’un plus grand nombre de veines croisées postnodales et postsubnodales, la base d’IR1 ayant plus de cellules éloignées de la base de RP2, un perostigma hyalin et un plus long RP3/4. Notre découverte augmente la diversité des demoiselles au cours du Crétacé moyen.</p>
         </trans-abstract>
         <kwd-group>
            <unstructured-kwd-group>Platycnemididae, Zygoptera, Odonata, Cenomanian, Cretaceous, Burmese amber</unstructured-kwd-group>
         </kwd-group>
         <kwd-group xml:lang="fr">
            <unstructured-kwd-group>Platycnemididae, Zygoptera, Odonates, Cénomanien, Crétacé, Ambre birman</unstructured-kwd-group>
         </kwd-group>
         <custom-meta-group>
            <custom-meta>
               <meta-name>presented</meta-name>
               <meta-value>Handled by Annalisa Ferretti</meta-value>
            </custom-meta>
         </custom-meta-group>
      </article-meta>
   </front>
   <body>
      <sec id="sec0005">
         <label>1</label>
         <title id="sect0025">Introduction</title>
         <p id="par0005">Platycnemididae <xref rid="bib0105" ref-type="bibr">Yakobson and Bianki, 1905</xref>, currently consists of over 400 species and is widely distributed in the Old World (<xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref>, <xref rid="bib0055" ref-type="bibr">Orr and Kalkman, 2010</xref>, <xref rid="bib0085" ref-type="bibr">Schorr and Paulson, 2015</xref> and <xref rid="bib0100" ref-type="bibr">Theischinger et al., 2015</xref>). Adults of this family are characterized by a laterally expanded head and the tibiae bearing dense long spines (<xref rid="bib0015" ref-type="bibr">Carle et al., 2008</xref>, <xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref> and <xref rid="bib0065" ref-type="bibr">Rehn, 2003</xref>). Platycnemididae is divided into six extant subfamilies after a recent molecular phylogeny of <xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref>: Allocnemidinae <xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref>, Calicnemiinae <xref rid="bib0035" ref-type="bibr">Fraser, 1957</xref>, Disparoneurinae <xref rid="bib0035" ref-type="bibr">Fraser, 1957</xref>, Idiocnemidinae <xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref>, Onychargiinae <xref rid="bib0025" ref-type="bibr">Dijkstra et al., 2014</xref>, and Platycnemidinae <xref rid="bib0105" ref-type="bibr">Yakobson and Bianki, 1905</xref>. A fossil subfamily Palaeodisparoneurinae Poinar, Bechly et Buckley, 2010 (type species: <italic>Palaeodisparoneura burmanica</italic> Poinar, Bechly et Buckley, 2010) was attributed to Platycnemididae based on a well-preserved damselfly from mid-Cretaceous Burmese amber; however, this subfamily was out of consideration of the molecular results provided by <xref rid="bib0025" ref-type="bibr">Dijkstra et al. (2014)</xref>. <italic>P. burmanica</italic> cannot be attributed to any recent taxa since it possesses the following unique autapomorphies: the midfork (the base of RP3/4) originates midway between the arculus and nodus, the pterostigma is of rectangular shape, RP1 is strongly kinked at the pterostigmal brace, and IR1 originates below the pterostigma. It is attributed to Platycnemididae by the presence of a special shape of the short male terminalia (superior appendages shaped like the hammer of a revolver), which is shared by Disparoneurinae and Caconeurinae (<xref rid="bib0060" ref-type="bibr">Poinar et al., 2010</xref>).</p>
         <p id="par0010">Here we describe a new species of <italic>Palaeodisparoneura</italic>, <italic>P. cretacica</italic> sp. nov., from Burmese amber.</p>
      </sec>
      <sec id="sec0010">
         <label>2</label>
         <title id="sect0030">Material and methods</title>
         <sec>
            <p id="par0015">The specimen described herein was collected from the Hukawng Valley of Kachin Province, Myanmar (locality in <xref rid="bib0045" ref-type="bibr">Kania et al., 2015</xref>: fig. 1). The age of Burmese amber is radiometrically dated at 98.79 ± 0.62 Ma (Earliest Cenomanian; <xref rid="bib0020" ref-type="bibr">Cohen et al., 2013</xref>) based on U–Pb zircon dating of the volcanoclastic matrix (<xref rid="bib0095" ref-type="bibr">Shi et al., 2012</xref>).</p>
         </sec>
         <sec>
            <p id="par0020">The amber containing the damselfly is yellow and transparent. Photographs were taken using a Zeiss Stereo Discovery V16 microscope system and Zen software. In most instances, incident and transmitted light were used simultaneously. All images are digitally stacked photomicrographic composites of approximately 40 individual focal planes obtained using the free software Combine ZP for a better illustration of the 3D structures. The line drawings were prepared from photographs using image-editing software (CorelDraw X7 and Adobe Photoshop CS6). The specimen is housed in the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences (NIGPAS).</p>
         </sec>
         <sec>
            <p id="par0025">The nomenclature of the dragonfly wing venation used in this paper is based on the interpretations of <xref rid="bib0070" ref-type="bibr">Riek (1976)</xref> and <xref rid="bib0075" ref-type="bibr">Riek and Kukalová-Peck (1984)</xref>, as modified by <xref rid="bib0050" ref-type="bibr">Nel et al. (1993)</xref> and <xref rid="bib0005" ref-type="bibr">Bechly (1996)</xref>. The higher classification of fossil and extant Odonatoptera, as well as family and generic characters followed in the present work, are based on the phylogenetic system proposed by <xref rid="bib0005" ref-type="bibr">Bechly (1996)</xref> and <xref rid="bib0025" ref-type="bibr">Dijkstra et al. (2014)</xref> for the phylogeny of extant Zygoptera. Wing abbreviations are as follows: Cr, nodal crossvein; CuA, cubitus anterior; DC, discoidal cell; IR, intercalary radial veins; MA, median anterior; MP, median posterior; N, nodus; Pt, pterostigma; RA, radius anterior; RP, radius posterior; Sn, subnodal crossvein. All measurements are given in mm.</p>
         </sec>
      </sec>
      <sec id="sec0015">
         <label>3</label>
         <title id="sect0035">Systematic palaeontology</title>
         <sec>
            <p id="par0030">Order Odonata <xref rid="bib0030" ref-type="bibr">Fabricius, 1793</xref>
            </p>
         </sec>
         <sec>
            <p id="par0035">Suborder Zygoptera <xref rid="bib0090" ref-type="bibr">Selys-Longchamps, 1854</xref>
            </p>
         </sec>
         <sec>
            <p id="par0040">Family Platycnemididae Yakobson and Bianki, 1905</p>
         </sec>
         <sec>
            <p id="par0045">Subfamily Palaeodisparoneurinae Poinar, Bechly et Buckley, 2010</p>
         </sec>
         <sec>
            <p id="par0050">Genus: <italic>
                  <bold>Palaeodisparoneura</bold>
               </italic> Poinar, Bechly et Buckley, 2010</p>
         </sec>
         <sec>
            <p id="par0055">
               <bold>Type species</bold>. <italic>Palaeodisparoneura burmanica</italic> Poinar, Bechly et Buckley, 2010</p>
         </sec>
         <sec>
            <p id="par0060">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov. (<xref rid="fig0005" ref-type="fig">Fig. 1</xref>, <xref rid="fig0010" ref-type="fig">Fig. 2</xref>, <xref rid="fig0015" ref-type="fig">Fig. 3</xref>, <xref rid="fig0020" ref-type="fig">Fig. 4</xref>, <xref rid="fig0025" ref-type="fig">Fig. 5</xref> and <xref rid="fig0030" ref-type="fig">Fig. 6</xref>)</p>
         </sec>
         <sec>
            <p id="par0065">
               <bold>Etymology</bold>
               <italic>.</italic> After the age of the species.</p>
         </sec>
         <sec>
            <p id="par0070">
               <bold>Material</bold>. Holotype. NIGP164075, four wings associated with a fragmentary abdomen and fragmentary legs, gender feminine; deposited in the Nanjing Institute of Geology and Palaeontology, Chinese Academy of Sciences, China.</p>
         </sec>
         <sec>
            <p id="par0075">
               <bold>Type stratum and locality</bold>. Hukawng Valley, Kachin Province, Myanmar; Lowermost Cenomanian, lowermost Upper Cretaceous.</p>
         </sec>
         <sec>
            <p id="par0080">
               <bold>Diagnosis</bold>
               <italic>.</italic> Eight postnodal crossveins and eight postsubnodal crossveins present before Pt, well aligned; IR1 seven or eight cells distal of base of RP2; RP3/4 long, ending on posterior wing margin two or three cells distal of base of IR1; Pt hyaline; tarsi armed with dense spines.</p>
         </sec>
         <sec>
            <p id="par0085">
               <bold>Description</bold>. All wings with basal part somewhat absent. Abdomen with basal five segments preserved, 13.39 mm long; gender feminine in absence of male accessory genitalia on ventral side of segment 2. Right forewing most completed preserved (<xref rid="fig0010" ref-type="fig">Figs. 2B, 3</xref>). Wing length 7.88 mm, maximum width 2.25 mm; length from base of RP3/4 to Pt base 5.61 mm, from Pt base to wing apex 2.03 mm. Eight postnodal crossveins and eight postsubnodal crossveins present before Pt, well aligned. Four postnodal crossveins and four postsubnodal crossveins present distal of Pt, non-aligned. Nodal structures not well preserved; Sn present and aligned with base of IR2. Midfork one long cell and 0.66 mm basal of Sn, ending on posterior margin three cells distal of base of IR1. IR2 basally straight but distally zigzagged. Base of RP2 three cells distal of Sn, lying 1.28 mm distally, nearer to N than to Pt. Base of IR1 eight cells distal of base of RP2, lying 3.81 mm distally and originating just below Pt. RP1 with a strong angle below pterostigmal brace. MA distally zigzagged and long, reaching posterior wing margin just below Pt-brace. MP curved and short. Pt one cell long, rectangular and hyaline (<xref rid="fig0025" ref-type="fig">Fig. 5</xref>), 0.64 mm long and 0.21 mm wide, strongly braced; Pt-brace more oblique than base of Pt. All intercalary veins (except for IR1 and IR2) suppressed. Longitudinal veins RA, IR1, RP1, IR2 and RP2 strongly converging to wing apex. Other wings resemble this wing except for following differences: IR1 seven cells distal of base of RP2 in left hindwing (<xref rid="fig0010" ref-type="fig">Fig. 2</xref>C); N well preserved (<xref rid="fig0020" ref-type="fig">Fig. 4</xref>) in left forewing, and Sn aligned with base of IR2; MP one cell length distal of CuA, ending on posterior wing margin just below N in left hindwing (<xref rid="fig0020" ref-type="fig">Fig. 4</xref>); CuA short and oblique in left hindwing (<xref rid="fig0020" ref-type="fig">Fig. 4</xref>); distal part of DC and subdiscoidal cell rectangular in left hindwing (<xref rid="fig0020" ref-type="fig">Fig. 4</xref>). Four fragmentary legs preserved (<xref rid="fig0030" ref-type="fig">Fig. 6</xref>), paired long spines present on tibia and tarsi; tarsi armed with about 14 pairs of spines.</p>
         </sec>
      </sec>
      <sec id="sec0020">
         <label>4</label>
         <title id="sect0040">Discussion</title>
         <sec>
            <p id="par0090">
               <italic>P.</italic> <italic>cretacica</italic> can be attributed to the subfamily Palaeodisparoneurinae Poinar, Bechly et Buckley, 2010 by sharing the characters provided by <xref rid="bib0060" ref-type="bibr">Poinar et al. (2010)</xref>: small damselfly species characterized by greatly shortened veins MP (only one cell long) and CuA (only an oblique crossvein); aligned postnodal and postsubnodal crossveins; rectangular and distinctly braced Pt; RP1 strongly kinked at pterostigmal brace; very short IR1 originating beneath Pt. <italic>Palaeodisparoneura</italic> Poinar, Bechly et Buckley, 2010 is also characterized by a rectangular discoidal cell, a well-developed sub-rectangular subdiscoidal cell being not fused to the wing margin, the midfork being situated halfway between the arculus and subnodus, and the male terminalia being short. These characters are poorly discernible in <italic>P. cretacica</italic>. However, the rectangular distal parts of the discoidal cell and subdiscoidal cell further support the attribution.</p>
         </sec>
         <sec>
            <p id="par0095">Although <italic>P. cretacica</italic> shares the diagnosis of <italic>P. burmanica</italic>, some obvious differences can be observed between these two species. Specifically, <italic>P. cretacica</italic> has more postnodal crossveins and postsubnodal crossveins (eight instead of five in <italic>P. burmanica</italic>) before the pterostigma, the base of IR1 is seven or eight cells distal of the base of RP2 instead of five rows in <italic>P. burmanica</italic>, the pterostigma is hyaline instead of dark colour, a long RP3/4 ends on the posterior wing margin three cells distal of the base of IR1 instead of one cell in <italic>P. burmanica</italic>. In view of the above differences, we suggest a new species for the new specimen.</p>
         </sec>
         <sec>
            <p id="par0100">The recently described odonatans in Burmese amber help us understand the role of Burma in odonatan diversification during the mid-Cretaceous. The damselfly group is represented by some extant families, viz., Platycnemididae, Platystictidae, Hemiphlebiidae and Perilestidae (<xref rid="bib0040" ref-type="bibr">Huang et al., 2015</xref>, <xref rid="bib0060" ref-type="bibr">Poinar et al., 2010</xref> and <xref rid="bib0110" ref-type="bibr">Zheng et al., 2016a</xref>, b, c). Besides <italic>P. burmanica</italic>, <italic>Cretadisparoneura hongi</italic> Huang, Azar, Cai et Nel, 2015 is the third representative of Platycnemididae (<xref rid="bib0040" ref-type="bibr">Huang et al., 2015</xref>). <italic>Burmalindenia imperfecta</italic>
               <xref rid="bib0080" ref-type="bibr">Schädel and Bechly, 2016</xref> is the first true dragonfly from Burmese amber (<xref rid="bib0080" ref-type="bibr">Schädel and Bechly, 2016</xref>). <italic>Burmaphlebia reifi</italic>
               <xref rid="bib0010" ref-type="bibr">Bechly and Poinar, 2013</xref> is the first damsel-dragonfly from amber (<xref rid="bib0010" ref-type="bibr">Bechly and Poinar, 2013</xref>). <italic>Burmahemiphlebia zhangi</italic>
               <xref rid="bib0110" ref-type="bibr">Zheng et al., 2016a</xref> is the first member of the Hemiphlebiidae in Burmese amber and may be the most common damselfly in these deposits, since about 40 specimens have been observed by the present authors (<xref rid="bib0110" ref-type="bibr">Zheng et al., 2016a</xref>). <italic>Palaeodysagrion cretacica</italic>
               <xref rid="bib0110" ref-type="bibr">Zheng et al., 2016a</xref> represents the second Cretaceous dysagrionid damselfly (<xref rid="bib0110" ref-type="bibr">Zheng et al., 2016a</xref>). <italic>Mesosticta burmatica</italic> Huang, Azar, Cai et Nel, 2015 and <italic>Mesosticta electronica</italic>
               <xref rid="bib0115" ref-type="bibr">Zheng et al., 2016b</xref> are the earliest fossil platystictid damselflies, which puts the origin of Platystictidae to at least mid-Cretaceous (<xref rid="bib0040" ref-type="bibr">Huang et al., 2015</xref> and <xref rid="bib0115" ref-type="bibr">Zheng et al., 2016b</xref>). <italic>Palaeoperilestes electronicus</italic>
               <xref rid="bib0120" ref-type="bibr">Zheng et al., 2016c</xref> is the first fossil representative of perilestid damselfly (<xref rid="bib0120" ref-type="bibr">Zheng et al., 2016c</xref>).</p>
         </sec>
      </sec>
      <sec id="sec0025">
         <label>5</label>
         <title id="sect0045">Conclusions</title>
         <sec>
            <p id="par0105">Abundant extant damselfly families have been discovered from mid-Cretaceous Burmese amber, indicating the important geological position of these tropical forests for early damselfly diversification. In the present paper, a new platycnemid damselfly, <italic>P. cretacica</italic> sp. nov., representing the second species of the extinct subfamily Palaeodisparoneurinae Poinar, Bechly et Buckley, 2010, is described from Burmese amber.</p>
         </sec>
      </sec>
   </body>
   <back>
      <ack>
         <title id="sect0050">Acknowledgements</title>
         <p id="par0110">This research was supported (grant numbers in parentheses) by the <funding-source id="gs1">
               <institution-wrap>
                  <institution>National Basic Research Program of China</institution>
               </institution-wrap>
            </funding-source> (<award-id award-type="grant" rid="gs1">2012CB821900</award-id>), <funding-source id="gs2">
               <institution-wrap>
                  <institution>National Natural Science Foundation of China</institution>
                  <institution-id>http://dx.doi.org/10.13039/501100001809</institution-id>
               </institution-wrap>
            </funding-source> (<award-id award-type="grant" rid="gs2">41572010, J1210006</award-id>), <funding-source id="gs3">
               <institution-wrap>
                  <institution>Youth Innovation Promotion Association of Chinese Academy of Sciences</institution>
                  <institution-id>http://dx.doi.org/10.13039/501100004739</institution-id>
               </institution-wrap>
            </funding-source> (<award-id award-type="grant" rid="gs3">2011224</award-id>), and the <funding-source id="gs4">
               <institution-wrap>
                  <institution>HKU Seed Funding Program for Basic Research</institution>
                  <institution-id>http://dx.doi.org/10.13039/100007916</institution-id>
               </institution-wrap>
            </funding-source> (<award-id award-type="grant" rid="gs4">201210159058</award-id>). We offer our sincere gratitude to Sandrine Fléchel and Dr. Annalisa Ferretti for editing, and two anonymous reviewers for the very useful comments on the earlier version of the manuscript.</p>
      </ack>
      <ref-list>
         <ref id="bib0005">
            <label>Bechly, 1996</label>
            <element-citation id="sbref0005" publication-type="article">
               <name>
                  <surname>Bechly</surname>
                  <given-names>G.</given-names>
               </name>
               <article-title>Morphologische Untersuchungen am Flügelgeäder der rezenten Libellen und deren Stammgruppenvertreter (Insecta; Pterygota; Odonata), unter besonderer Berücksichtigung der Phylogenetischen Systematik und des Grundplanes der Odonata</article-title>
               <source>Petalura</source>
               <volume>2</volume>
               <year>1996</year>
               <page-range>1–402</page-range>
            </element-citation>
         </ref>
         <ref id="bib0010">
            <label>Bechly and Poinar, 2013</label>
            <element-citation id="sbref0010" publication-type="article">
               <name>
                  <surname>Bechly</surname>
                  <given-names>G.</given-names>
               </name>
               <name>
                  <surname>Poinar</surname>
                  <given-names>G.</given-names>
                  <suffix>Jr.</suffix>
               </name>
               <article-title>
                  <italic>Burmaphlebia reifi</italic> gen. et sp. nov., the first anisozygopteran damsel-dragonfly (Odonata: Epiophlebioptera: Burmaphlebiidae fam. nov.) from Early Cretaceous Burmese amber</article-title>
               <source>Hist. Biol.</source>
               <volume>25</volume>
               <year>2013</year>
               <page-range>233–237</page-range>
            </element-citation>
         </ref>
         <ref id="bib0015">
            <label>Carle et al., 2008</label>
            <element-citation id="sbref0015" publication-type="article">
               <name>
                  <surname>Carle</surname>
                  <given-names>F.L.</given-names>
               </name>
               <name>
                  <surname>Kjer</surname>
                  <given-names>K.M.</given-names>
               </name>
               <name>
                  <surname>May</surname>
                  <given-names>M.L.</given-names>
               </name>
               <article-title>Evolution of Odonata, with special reference to Coenagrionoidea (Zygoptera)</article-title>
               <source>Arthropod Syst. Phylo.</source>
               <volume>66</volume>
               <year>2008</year>
               <page-range>37–44</page-range>
            </element-citation>
         </ref>
         <ref id="bib0020">
            <label>Cohen et al., 2013</label>
            <element-citation id="sbref0020" publication-type="article">
               <name>
                  <surname>Cohen</surname>
                  <given-names>K.M.</given-names>
               </name>
               <name>
                  <surname>Finney</surname>
                  <given-names>S.C.</given-names>
               </name>
               <name>
                  <surname>Gibbard</surname>
                  <given-names>P.L.</given-names>
               </name>
               <name>
                  <surname>Fan</surname>
                  <given-names>J.</given-names>
               </name>
               <article-title>The ICS international chronostratigraphic chart</article-title>
               <source>Episodes</source>
               <volume>36</volume>
               <year>2013</year>
               <page-range>199–204</page-range>
            </element-citation>
         </ref>
         <ref id="bib0025">
            <label>Dijkstra et al., 2014</label>
            <element-citation id="sbref0025" publication-type="article">
               <name>
                  <surname>Dijkstra</surname>
                  <given-names>K.-D.B.</given-names>
               </name>
               <name>
                  <surname>Kalkman</surname>
                  <given-names>V.J.</given-names>
               </name>
               <name>
                  <surname>Dow</surname>
                  <given-names>R.A.</given-names>
               </name>
               <name>
                  <surname>Stokvis</surname>
                  <given-names>F.R.</given-names>
               </name>
               <name>
                  <surname>van Tol</surname>
                  <given-names>J.</given-names>
               </name>
               <article-title>Redefining the damselfly families: a comprehensive molecular phylogeny of Zygoptera (Odonata)</article-title>
               <source>Syst. Entomol.</source>
               <volume>39</volume>
               <year>2014</year>
               <page-range>68–96</page-range>
            </element-citation>
         </ref>
         <ref id="bib0030">
            <label>Fabricius, 1793</label>
            <element-citation id="sbref0030" publication-type="article">
               <name>
                  <surname>Fabricius</surname>
                  <given-names>J.C.</given-names>
               </name>
               <article-title>Entomologia systematica emendata et aucta, secundum classes, ordines, genera, species, adjectis synonymis, locis, observationibus, descriptionibus</article-title>
               <source>C. G. Proft, Hafniae (= Copenhagen)</source>
               <volume>3</volume>
               <year>1793</year>
               <comment>1–487 and 1–349</comment>
            </element-citation>
         </ref>
         <ref id="bib0035">
            <label>Fraser, 1957</label>
            <element-citation id="sbref0035" publication-type="article">
               <name>
                  <surname>Fraser</surname>
                  <given-names>F.C.</given-names>
               </name>
               <article-title>A reclassification of the order Odonata, based on some new interpretations of the venation of the dragonfly wing</article-title>
               <source>R. Zool. Soc. N. S. W. Sydney</source>
               <volume>12</volume>
               <year>1957</year>
               <page-range>1–133</page-range>
            </element-citation>
         </ref>
         <ref id="bib0040">
            <label>Huang et al., 2015</label>
            <element-citation id="sbref0040" publication-type="article">
               <name>
                  <surname>Huang</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Azar</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Cai</surname>
                  <given-names>C.</given-names>
               </name>
               <name>
                  <surname>Nel</surname>
                  <given-names>A.</given-names>
               </name>
               <article-title>New damselfly genera in the Cretaceous Burmese amber attributable to the Platystictidae and Platycnemididae Disparoneurinae (Odonata: Zygoptera)</article-title>
               <source>Cretaceous Res.</source>
               <volume>56</volume>
               <year>2015</year>
               <page-range>237–243</page-range>
            </element-citation>
         </ref>
         <ref id="bib0045">
            <label>Kania et al., 2015</label>
            <element-citation id="sbref0045" publication-type="article">
               <name>
                  <surname>Kania</surname>
                  <given-names>I.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>B.</given-names>
               </name>
               <name>
                  <surname>Szwedo</surname>
                  <given-names>J.</given-names>
               </name>
               <article-title>
                  <italic>Dicranoptycha</italic> Osten Sacken, 1860 (Diptera, Limoniidae) from the earliest Upper Cretaceous Burmese amber</article-title>
               <source>Cretaceous Res.</source>
               <volume>52</volume>
               <year>2015</year>
               <page-range>522–530</page-range>
            </element-citation>
         </ref>
         <ref id="bib0050">
            <label>Nel et al., 1993</label>
            <element-citation id="sbref0050" publication-type="article">
               <name>
                  <surname>Nel</surname>
                  <given-names>A.</given-names>
               </name>
               <name>
                  <surname>Martinez-Delclòs</surname>
                  <given-names>X.</given-names>
               </name>
               <name>
                  <surname>Paicheler</surname>
                  <given-names>J.C.</given-names>
               </name>
               <name>
                  <surname>Henrotay</surname>
                  <given-names>M.</given-names>
               </name>
               <article-title>Les ‘Anisozygoptera’ fossiles. Phylogénie et classification (Odonata)</article-title>
               <source>Martinia Num. Hors Sér.</source>
               <volume>3</volume>
               <year>1993</year>
               <page-range>1–311</page-range>
            </element-citation>
         </ref>
         <ref id="bib0055">
            <label>Orr and Kalkman, 2010</label>
            <element-citation id="sbref0055" publication-type="article">
               <name>
                  <surname>Orr</surname>
                  <given-names>A.G.</given-names>
               </name>
               <name>
                  <surname>Kalkman</surname>
                  <given-names>V.J.</given-names>
               </name>
               <article-title>
                  <italic>Arrhenocnemis parvibullis</italic> sp. nov. (Odonata: Platycnemididae), a new calicnemiine damselfly from Papua New Guinea, with a description of the female of A. amphidactylis Lieftinck, 1949</article-title>
               <source>Aust. Entomol.</source>
               <volume>37</volume>
               <issue>4</issue>
               <year>2010</year>
               <page-range>137–146</page-range>
            </element-citation>
         </ref>
         <ref id="bib0060">
            <label>Poinar et al., 2010</label>
            <element-citation id="sbref0060" publication-type="article">
               <name>
                  <surname>Poinar</surname>
                  <given-names>G.</given-names>
                  <suffix>Jr.</suffix>
               </name>
               <name>
                  <surname>Bechly</surname>
                  <given-names>G.</given-names>
               </name>
               <name>
                  <surname>Buckley</surname>
                  <given-names>R.</given-names>
               </name>
               <article-title>First record of Odonata and a new subfamily of damselflies from Early Cretaceous Burmese amber</article-title>
               <source>Palaeodiversity</source>
               <volume>3</volume>
               <year>2010</year>
               <page-range>15–22</page-range>
            </element-citation>
         </ref>
         <ref id="bib0065">
            <label>Rehn, 2003</label>
            <element-citation id="sbref0065" publication-type="article">
               <name>
                  <surname>Rehn</surname>
                  <given-names>A.C.</given-names>
               </name>
               <article-title>Phylogenetic analysis of higher-level relationships of Odonata</article-title>
               <source>Syst. Entomol.</source>
               <volume>28</volume>
               <year>2003</year>
               <page-range>181–239</page-range>
            </element-citation>
         </ref>
         <ref id="bib0070">
            <label>Riek, 1976</label>
            <element-citation id="sbref0070" publication-type="article">
               <name>
                  <surname>Riek</surname>
                  <given-names>E.F.</given-names>
               </name>
               <article-title>A new collection of insects from the Upper Triassic of South Africa</article-title>
               <source>Ann. Natal Mus.</source>
               <volume>22</volume>
               <year>1976</year>
               <page-range>791–820</page-range>
            </element-citation>
         </ref>
         <ref id="bib0075">
            <label>Riek and Kukalová-Peck, 1984</label>
            <element-citation id="sbref0075" publication-type="article">
               <name>
                  <surname>Riek</surname>
                  <given-names>E.F.</given-names>
               </name>
               <name>
                  <surname>Kukalová-Peck</surname>
                  <given-names>J.</given-names>
               </name>
               <article-title>A new interpretation of dragonfly wing venation based upon Early Carboniferous fossils from Argentina (Insecta: Odonatoidea) and basic character states in pterygote wings</article-title>
               <source>Can. J. Zool.</source>
               <volume>62</volume>
               <year>1984</year>
               <page-range>1150–1166</page-range>
            </element-citation>
         </ref>
         <ref id="bib0080">
            <label>Schädel and Bechly, 2016</label>
            <element-citation id="sbref0080" publication-type="article">
               <name>
                  <surname>Schädel</surname>
                  <given-names>M.</given-names>
               </name>
               <name>
                  <surname>Bechly</surname>
                  <given-names>G.</given-names>
               </name>
               <article-title>First record of Anisoptera (Insecta: Odonata) from mid-Cretaceous Burmese Amber</article-title>
               <source>Zootaxa</source>
               <volume>4103</volume>
               <issue>6</issue>
               <year>2016</year>
               <page-range>537–549</page-range>
            </element-citation>
         </ref>
         <ref id="bib0085">
            <label>Schorr and Paulson, 2015</label>
            <element-citation id="sbref0085" publication-type="book">
               <name>
                  <surname>Schorr</surname>
                  <given-names>M.</given-names>
               </name>
               <name>
                  <surname>Paulson</surname>
                  <given-names>D.</given-names>
               </name>
               <source>World Odonata list</source>
               <year>2015</year>
               <comment>Available on the Internet at: <ext-link xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="http://www.pugetsound.edu/">http://www.pugetsound.edu/</ext-link> (accessed 16 March 2016)</comment>
            </element-citation>
         </ref>
         <ref id="bib0090">
            <label>Selys-Longchamps, 1854</label>
            <element-citation id="sbref0090" publication-type="article">
               <name>
                  <surname>Selys-Longchamps</surname>
                  <given-names>E.</given-names>
                  <suffix>de</suffix>
               </name>
               <article-title>Monographie des Caloptérygines</article-title>
               <source>Mémoires de la Société royale des Sciences de Liège</source>
               <volume>9</volume>
               <year>1854</year>
               <comment>xi + 291 pp</comment>
            </element-citation>
         </ref>
         <ref id="bib0095">
            <label>Shi et al., 2012</label>
            <element-citation id="sbref0095" publication-type="article">
               <name>
                  <surname>Shi</surname>
                  <given-names>G.</given-names>
               </name>
               <name>
                  <surname>Grimaldi</surname>
                  <given-names>D.A.</given-names>
               </name>
               <name>
                  <surname>Harlow</surname>
                  <given-names>G.E.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>J.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>J.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>M.</given-names>
               </name>
               <name>
                  <surname>Lei</surname>
                  <given-names>W.</given-names>
               </name>
               <name>
                  <surname>Li</surname>
                  <given-names>Q.</given-names>
               </name>
               <name>
                  <surname>Li</surname>
                  <given-names>X.</given-names>
               </name>
               <article-title>Age constraint on Burmese amber based on U-Pb dating of zircons</article-title>
               <source>Cretaceous Res.</source>
               <volume>37</volume>
               <year>2012</year>
               <page-range>155–163</page-range>
            </element-citation>
         </ref>
         <ref id="bib0100">
            <label>Theischinger et al., 2015</label>
            <element-citation id="sbref0100" publication-type="article">
               <name>
                  <surname>Theischinger</surname>
                  <given-names>G.</given-names>
               </name>
               <name>
                  <surname>Gassmann</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Richards</surname>
                  <given-names>S.J.</given-names>
               </name>
               <article-title>
                  <italic>Macrocnemis gracilis</italic>, a new genus and species of Idiocnemidinae (Zygoptera: Platycnemididae) from Papua New Guinea</article-title>
               <source>Zootaxa</source>
               <volume>3990</volume>
               <issue>3</issue>
               <year>2015</year>
               <page-range>429–437</page-range>
            </element-citation>
         </ref>
         <ref id="bib0105">
            <label>Yakobson and Bianki, 1905</label>
            <element-citation id="sbref0105" publication-type="book">
               <name>
                  <surname>Yakobson</surname>
                  <given-names>G.G.</given-names>
               </name>
               <name>
                  <surname>Bianki</surname>
                  <given-names>V.L.</given-names>
               </name>
               <source>Orthoptera and Pseudo-Neuroptera of the Russian Empire and adjoining countries</source>
               <year>1905</year>
               <publisher-name>St. Petersburg</publisher-name>
               <comment>(In Russian)</comment>
            </element-citation>
         </ref>
         <ref id="bib0110">
            <label>Zheng et al., 2016a</label>
            <element-citation id="sbref0110" publication-type="book">
               <name>
                  <surname>Zheng</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Zhang</surname>
                  <given-names>Q.</given-names>
               </name>
               <name>
                  <surname>Nel</surname>
                  <given-names>A.</given-names>
               </name>
               <name>
                  <surname>Jarzembowski</surname>
                  <given-names>E.A.</given-names>
               </name>
               <name>
                  <surname>Zhou</surname>
                  <given-names>Z.</given-names>
               </name>
               <name>
                  <surname>Chang</surname>
                  <given-names>S.-C.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>B.</given-names>
               </name>
               <source>New damselflies (Odonata: Zygoptera: Hemiphlebiidae, Dysagrionidae) from mid-Cretaceous Burmese amber.</source>
               <year>2016</year>
               <publisher-name>Alcheringa</publisher-name>
               <pub-id pub-id-type="doi">10.1080/03115518.2016.1164402</pub-id>
            </element-citation>
         </ref>
         <ref id="bib0115">
            <label>Zheng et al., 2016b</label>
            <element-citation id="sbref0115" publication-type="article">
               <name>
                  <surname>Zheng</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Zhang</surname>
                  <given-names>Q.</given-names>
               </name>
               <name>
                  <surname>Chang</surname>
                  <given-names>S.-C.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>B.</given-names>
               </name>
               <article-title>A new damselfly (Odonata: Zygoptera: Platystictidae) from mid-Cretaceous Burmese amber</article-title>
               <source>Cretaceous Res.</source>
               <volume>63</volume>
               <year>2016</year>
               <page-range>142–147</page-range>
            </element-citation>
         </ref>
         <ref id="bib0120">
            <label>Zheng et al., 2016c</label>
            <element-citation id="sbref0120" publication-type="article">
               <name>
                  <surname>Zheng</surname>
                  <given-names>D.</given-names>
               </name>
               <name>
                  <surname>Wang</surname>
                  <given-names>B.</given-names>
               </name>
               <name>
                  <surname>Jarzembowski</surname>
                  <given-names>E.A.</given-names>
               </name>
               <name>
                  <surname>Chang</surname>
                  <given-names>S.-C.</given-names>
               </name>
               <name>
                  <surname>Nel</surname>
                  <given-names>A.</given-names>
               </name>
               <article-title>The first fossil Perilestidae (Odonata: Zygoptera) from mid-Cretaceous Burmese amber</article-title>
               <source>Cretaceous Res.</source>
               <volume>65</volume>
               <year>2016</year>
               <page-range>199–205</page-range>
            </element-citation>
         </ref>
      </ref-list>
   </back>
   <floats-group>
      <fig id="fig0005">
         <label>Fig. 1</label>
         <caption>
            <p id="spar0015">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075, photomicrograph of specimen.</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0020">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov., holotype, NIGP164075, microphotographie de l’échantillon.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr1.jpg"/>
      </fig>
      <fig id="fig0010">
         <label>Fig. 2</label>
         <caption>
            <p id="spar0025">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075. A: photomicrograph of right forewing; B: photomicrograph of right hindwing; C: photomicrograph of left forewing (upper) and hindwing (lower).</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0030">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov., holotype, NIGP164075. A : microphotographie de l’aile antérieure droite ; B : microphotographie de l’aile postérieure droite ; C : microphotographie de l’aile antérieure gauche (en haut) et de l’aile postérieure gauche (en bas).</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr2.jpg"/>
      </fig>
      <fig id="fig0015">
         <label>Fig. 3</label>
         <caption>
            <p id="spar0035">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075, line drawing showing right hindwing venation.</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0040">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov. holotype, NIGP164075, dessin au trait de la venation de l’aile postérieure droite.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr3.jpg"/>
      </fig>
      <fig id="fig0020">
         <label>Fig. 4</label>
         <caption>
            <p id="spar0045">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075, photomicrograph showing detail of basal wings.</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0050">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov., holotype, NIGP 164075, microphotographie montrant le détail des ailes basales.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr4.jpg"/>
      </fig>
      <fig id="fig0025">
         <label>Fig. 5</label>
         <caption>
            <p id="spar0055">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075, photomicrograph of right forewing pterostigma.</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0060">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov., holotype, NIGP164075, microphotographie du pterostigma de l’aile antérieure droite.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr5.jpg"/>
      </fig>
      <fig id="fig0030">
         <label>Fig. 6</label>
         <caption>
            <p id="spar0065">
               <italic>Palaeodisparoneura cretacica</italic> sp. nov., holotype, NIGP164075, photomicrograph of legs.</p>
         </caption>
         <caption xml:lang="fr">
            <p id="spar0070">
               <italic>Palaeodisparoneura cretacia</italic> sp. nov., holotype, NIGP164075, microphotographie des pattes.</p>
         </caption>
         <graphic xmlns:xlink="http://www.w3.org/1999/xlink" xlink:href="main.assets/gr6.jpg"/>
      </fig>
   </floats-group>
</article>