First Baltic amber megapodagrionid damselfly (Odonata: Zygoptera)

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<ul><li><p>This article was downloaded by: ["University at Buffalo Libraries"]On: 05 October 2014, At: 00:59Publisher: Taylor &amp; FrancisInforma Ltd Registered in England and Wales Registered Number: 1072954 Registered office: MortimerHouse, 37-41 Mortimer Street, London W1T 3JH, UK</p><p>Annales de la Socit entomologique de France(N.S.): International Journal of EntomologyPublication details, including instructions for authors and subscription information:http://www.tandfonline.com/loi/tase20</p><p>First Baltic amber megapodagrionid damselfly(Odonata: Zygoptera)Dany Azar a &amp; Andr Nel ba Department of Natural Sciences , Lebanese University , Fanar - Matn - P. O. box26110217 , Lebanonb Entomologie , CNRS UMR 5202, Museum National dHistoire Naturelle , CP 50, 45 rueBuffon, Paris , F-75005 , FrancePublished online: 31 May 2013.</p><p>To cite this article: Dany Azar &amp; Andr Nel (2008) First Baltic amber megapodagrionid damselfly (Odonata: Zygoptera),Annales de la Socit entomologique de France (N.S.): International Journal of Entomology, 44:4, 451-457, DOI:10.1080/00379271.2008.10697580</p><p>To link to this article: http://dx.doi.org/10.1080/00379271.2008.10697580</p><p>PLEASE SCROLL DOWN FOR ARTICLE</p><p>Taylor &amp; Francis makes every effort to ensure the accuracy of all the information (the Content) containedin the publications on our platform. However, Taylor &amp; Francis, our agents, and our licensors make norepresentations or warranties whatsoever as to the accuracy, completeness, or suitability for any purpose ofthe Content. Any opinions and views expressed in this publication are the opinions and views of the authors,and are not the views of or endorsed by Taylor &amp; Francis. The accuracy of the Content should not be reliedupon and should be independently verified with primary sources of information. Taylor and Francis shallnot be liable for any losses, actions, claims, proceedings, demands, costs, expenses, damages, and otherliabilities whatsoever or howsoever caused arising directly or indirectly in connection with, in relation to orarising out of the use of the Content.</p><p>This article may be used for research, teaching, and private study purposes. Any substantial or systematicreproduction, redistribution, reselling, loan, sub-licensing, systematic supply, or distribution in anyform to anyone is expressly forbidden. Terms &amp; Conditions of access and use can be found at http://www.tandfonline.com/page/terms-and-conditions</p><p>http://www.tandfonline.com/loi/tase20http://www.tandfonline.com/action/showCitFormats?doi=10.1080/00379271.2008.10697580http://dx.doi.org/10.1080/00379271.2008.10697580http://www.tandfonline.com/page/terms-and-conditionshttp://www.tandfonline.com/page/terms-and-conditions</p></li><li><p>Ann. soc. entomol. Fr. (n.s.), 2008, 44 (4) : 451-457</p><p>451</p><p>ARTICLE</p><p>First Baltic amber megapodagrionid damselfl y (Odonata: Zygoptera)</p><p>Abstract. Electropodagrion szwedoi n. gen., n. sp., fi rst Baltic amber megapodagrionid damselfl y, is described. The European and North American fossils document a very high diversity and a much wider distribution of this group of damselfl ies during the Cenozoic than today. A checklist of described fossil species of damselfl ies of the family Megapodagrionidae is given.</p><p>Rsum. Le premier Megapodagrionidae de lambre de la Baltique (Odonata : Zygoptera). Electropodagrion szwedoi n. gen., n. sp., premier Megapodagrionidae de lambre balte, est dcrit. Les fossiles europens et nord-amricains dmontrent une trs forte diversit et une distribution beaucoup plus large pour ce groupe pendant le Cnozoque que dans lactuel. Une liste des espces fossiles de Zygoptera: Megapodagrionidae est donne. Keywords: Fossil; Tertiary; Eocene; Baltic Amber.</p><p>Dany Azar (1) &amp; Andr Nel (2)*(1) Lebanese University / Faculty of Sciences II / Department of Natural Sciences/ Fanar - Matn - P. O. box 26110217 / Lebanon </p><p>(2) CNRS UMR 5202, Musum National dHistoire Naturelle, CP 50, Entomologie, 45 rue Buff on, F-75005, Paris, France * Corresponding author</p><p>E-mail: azar@mnhn.fr, anel@mnhn.frAccept le 23 octobre 2007</p><p>The damselfl y taxon Megapodagrionidae sensu Bechly 1996 currently includes 34 extant genera and shows an essentially pantropical distribution (Petruleviius et al. 2008). Relationships among the known living species, are however unclear, and it is unknown whether the fossil species are allied to those that are living today in the pantropical regions. Th e fossil record of the megapodagrionid damselfl ies indicates that this group have a long history extending well beyond the pantropical region.</p><p>Th e higher classifi cation in this work follows the phylogenetic system of fossil and extant Odonata of Bechly (1996, 2005). Megapodagrionid damselfl ies are classifi ed as Euzygoptera: Coenagriomorpha. In addition to Megapodagrionidae, the Coenagriomorpha comprises Hypolestidae and Coenagrioniformia in an unresolved phylogenetic relationship; their phylogenetic positions might be subject to change (sedis mutabilis) (Bechly 2005). Included in Megapodagrionidae are Argiolestinae and Megapodagrioninae. Until now, no synapomorphies for all Megapodagrionidae are known, so they could well be a paraphyletic group in the present composition (Bechly 1996, 2005; Rehn 2003).</p><p>We describe the fi rst megapodagrionid from the Baltic amber, which also represents the fi rst record of the family in amber.</p><p>Material and methodsTh e specimen was examined with an Olympus SZX9 stereomicroscope. Drawings were made using a camera lucida. All measurements were produced using an ocular micrometer; with structures measured as preserved. Photographs were made with an Olympus C5060 digital camera.Th e nomenclature of the dragonfl y wing venation is based on the interpretations of Riek (1976), Riek &amp; Kukalov-Peck (1984), amended by Kukalov-Peck (1991), Nel et al. (1993) and Bechly (1996).</p><p>Family Megapodagrionidae Calvert 1913</p><p>Genus Electropodagrion n. gen.Type species. Electropodagrion szwedoi n. sp.</p><p>Etymology. Named after Elektron, Greek name for amber and Podagrion.Diagnosis. Pterostigma short, covering one cell; pterostigmal brace weak and not oblique; no secondary longitudinal veins between main longitudinal veins except for two weak ones between IR2 and RP3/4 and RP3/4 and MA; CuP exactly opposite base of AA.</p><p>Electropodagrion szwedoi n. sp.(Figs 15)</p><p>Material. Holotype specimen number 4995, Museum Inkluzji W Bursztynie, Katedra Zoologii Bezkrgocw, Uniwersytet Gdaski, Poland. Etymology. Named after our friend and colleague Dr Jacek Szwedo.Description. Th e thorax with one leg, three wing bases and a distal two third of another wing are preserved. With all the </p><p>Dow</p><p>nloa</p><p>ded </p><p>by [</p><p>"Uni</p><p>vers</p><p>ity a</p><p>t Buf</p><p>falo</p><p> Lib</p><p>rari</p><p>es"]</p><p> at 0</p><p>0:59</p><p> 05 </p><p>Oct</p><p>ober</p><p> 201</p><p>4 </p></li><li><p>452</p><p>D. Azar &amp; A. Nel </p><p>preserved structures of the wings, we were able to reconstruct partially the forewing and entirely the hindwing (see fi gs 15). Wings nearly similar, hyaline, length 20 mm; width at nodus 3.3 mm; max. width (between nodus and pterostigma) 4.8 mm; distance from base to nodus 7.2 mm (nodus situated at about 36% of wing length); distance from nodus to pterostigma 10.1 mm; distance from base to arculus 4.1 mm; Ax1 and Ax2 well defi ned; Ax1 1.5 mm basal of arculus and Ax2 opposite arculus; no secondary antenodal cross-veins; nine postnodal cross-veins between nodus and pterostigma, rather well aligned with their corresponding postsubnodal cross-veins; pterostigma 1.5 mm long and max. 0.7 mm wide, covering one cell and a half; anterior and posterior margins of pterostigma not distinctly </p><p>broadened; basal side of pterostigma not oblique; fi ve cross-veins distal of pterostigma between C and RA; pterostigmal brace vein very slightly oblique; posterior part of arculus (= basal discoidal cross-vein) present, so discoidal cell basally closed, quadrangular, with distal side very oblique, and free of cross-veins, basal margin 0.17 mm long, costal margin 1.04 mm long, posterior margin 1.46 mm, distal margin 0.67 mm long; posterior margin of discoidal cell (= MP+CuA) exactly aligned with MP; origins of RP and MA strongly approximate in arculus; cubital cell free (except for CuP-crossing, 0.5 mm basal of arculus); anal area max. 0.25 mm wide with one row of cells; basal part of AA and AP fused, so that anal area not reaching wing base; CuP exactly opposite base of AA; cubito-</p><p>Figure 1 Electropodagrion szwedoi n. gen., n. sp., holotype 4995, photograph of forewing base.</p><p>Figure 2 Electropodagrion szwedoi n. gen., n. sp., holotype 4995, photograph of hind wing base.</p><p>Dow</p><p>nloa</p><p>ded </p><p>by [</p><p>"Uni</p><p>vers</p><p>ity a</p><p>t Buf</p><p>falo</p><p> Lib</p><p>rari</p><p>es"]</p><p> at 0</p><p>0:59</p><p> 05 </p><p>Oct</p><p>ober</p><p> 201</p><p>4 </p></li><li><p>First Baltic amber megapodagrionid</p><p>453</p><p>anal area max. 0.75 mm wide with one row of cells; CuA zigzagged; nodal Cr and subnodus very oblique; nodal Cr branching from ScP 0.2 mm basal of fusion of ScP with costal margin; MP, MA, RP3/4, and IR2 parallel; MP straight, not distally zigzagged, and ending between nodus and pterostigma; postdiscoidal area narrow; MA slightly zigzagged and ending two postnodal cells basal of pterostigma; RP3/4 and MA with one row of cells between them but with area distally widened; RP3/4 ending opposite basal side of pterostigma; base of RP3/4 0.9 mm basal of subnodus; IR2 aligned with subnodus; RP2 originating about three cells and 2.9 mm distal of subnodus; base of IR1 about three cells distally; no lestine oblique vein O between RP2 and IR2; RP2 and IR1, IR1 and RP1, and RP2 and IR2 with only one row of cells between them; RP1 without distinct kink at pterostigmal brace; RA and RP1 converging towards wing apex; wing base short.Th orax 2.2 mm wide; leg with femur 3.4 mm long, tibia 4.0 mm long, tarsus 1.5 mm long, femoral spines 0.7 mm long, tibial spines 1.1 mm long.</p><p>DiscussionElectropodagrion szwedoi n. gen., n. sp. can be </p><p>considered as a Coenagrionomorpha Bechly, 1996 because of the presence of the synapomorphies (sensu Bechly 1996) of this clade in its wing venation, i.e. pterostigma shortened; postnodal and postsubnodal </p><p>cross-veins aligned; lestine oblique vein absent; basal closure of discoidal cell including the development of a dorsal arcular bracket. Within the Coenagrionomorpha, the Hypolestidae can be excluded because of their amphipterygid type of pterostigma (basal margin strongly slanting). Th e Coenagrioniformia Bechly, 1996 can also be excluded because of their intercalary veins (except IR1 and IR2) suppressed and presence of only two rows of cells in the total wing space between RP1 and RP2; that are separated by the IR1. Th e last group within the Coenagriomorpha are the Megapodagrionidae. Th ey have no known synapomorphies in the wing venation (after Bechly 1996).</p><p>Rcenis (1959) proposed a division of the Megapodagrionidae into four subfamilies Argiolestinae Fraser 1957, Megapodagrioninae Calvert 1913, Philosininae Ris 1917, and Dysagrioninae Cockerell 1908 (transferred with the Th aumatoneurinae in Dysagrionidae, Bechly 1996; Rust et al. in press). Davies (1981) added the Tactonemidinae. Affi nities with this last subfamily are excluded because they have veins RP3/4 and IR2 arising distal of the </p><p>Figure 3 Electropodagrion szwedoi n. gen., n. sp., holotype 4995, photograph of hind wing mid part.</p><p>Dow</p><p>nloa</p><p>ded </p><p>by [</p><p>"Uni</p><p>vers</p><p>ity a</p><p>t Buf</p><p>falo</p><p> Lib</p><p>rari</p><p>es"]</p><p> at 0</p><p>0:59</p><p> 05 </p><p>Oct</p><p>ober</p><p> 201</p><p>4 </p></li><li><p>454</p><p>D. Azar &amp; A. Nel </p><p>subnodus. Electropodagrion shares with Philosina Ris, 1917, unique representative of the Philosininae, the structures of petiole and of anal and CuP veins, but it diff ers in its straight CuA with narrow cubito-anal area, distinctly shorter pterostigma, and absence of secondary longitudinal veins between RP1 and RP2 (Ris 1917).</p><p>An attribution to the Argiolestinae is unlikely because of the longitudinal wing veins straight in their </p><p>posterior part, and not shortened (especially MA, MP, and CuA), the branches of RP (RP3/4 and IR2) not distally diverging (Bechly 1996), and base of free AA basal of the arculus.</p><p>Th e Megapodagrioninae have no known synapomorphies in the wing venation sensu Bechly (1996). Only the representatives of the tribe Megapodagrionini Rcenis, 1959 have their veins CuP proximal of the arculus and distal of base of AA, </p><p>Figure 4 Electropodagrion szwedoi n. gen., n. sp., holotype 4995, photograph of hind wing apex.</p><p>Figure 5 Electropodagrion szwedoi n. gen., n. sp., holotype 4995, drawing of reconstruction fore- and hind wings (scale bar represents 0.5 mm).</p><p>Dow</p><p>nloa</p><p>ded </p><p>by [</p><p>"Uni</p><p>vers</p><p>ity a</p><p>t Buf</p><p>falo</p><p> Lib</p><p>rari</p><p>es"]</p><p> at 0</p><p>0:59</p><p> 05 </p><p>Oct</p><p>ober</p><p> 201</p><p>4 </p></li><li><p>First Baltic amber megapodagrionid</p><p>455</p><p>as in our fossil (see list in Nel et al. 1997). Among them, Priscagrion Zhou &amp; Wilson, 2001 has secondary antenodal cross-veins (Zhou &amp; Wilson 2001; De Marmels 2002). Megapodagrion Selys 1886 has a CuP in a more distal position relatively to base of AA, and its pterostigma is distinctly narrower than in Electropodagrion. Allopodagrion Frster 1910 and Teinopodagrion De Marmels 2001 have one or two secondary longitudinal veins between IR1 and RP2, unlike Electropodagrion (De Marmels 2001).</p><p>Rhinagrion Calvert 1913 has a pterostigma covering two or three cells, and a secondary longitudinal vein between MA and MP, unlike Electropodagrion. Mesopodagrion McLachlan 1896 has also a pterostigma covering three cells (McLachlan 1896).</p><p>Th e wing venation of the New Caledonian recent genus Trineuragrion Ris 1915 has strong similarities with that of Electropodagrion, especially in the short pterostigma, pterostigmal brace weak, CuP at base of AA, nearly no secondary longitudinal veins between main longitudinal veins. Th e few diff erences are as follows: Electropodagrion has less postnodal crossveins (nine instead of 14), base of RP2 in a more basal position (three cells distal of subnodus instead of 6-7) (Mnz 1919).</p><p>Nel et al. (1997) listed and commented the fossil megapodagrionid genera. An updated checklist of fossil Megapodagrionidae is given below (Appendix 1). Among them, Electropodagrion diff ers from Eckfeldia Petruleviius et al. (2008), in its pterostigma covering one cell. Furagrion Petruleviius et al. 2008 has its CuP in a more distal position, a pterostigma covering 23 cells, and secondary longitudinal veins between IR1 and RP2. Th e Megapodagrionidae genus and species undetermined from the Eocene of Messel (Germany) diff ers from Electropodagrion in the same two last characters (Petruleviius et al. 2008). Electropodagrion diff ers from Eopodagrion Cockerell 1920 in its pterostigmal brace not oblique (Cockerell 1920). Melanagrion Cockerell 1907 has two rows of cells in the cubito-anal area and in the area between C and RA distal of pterostigma, and its wing is petiolated distal of CuP. Miopodagrion Kennedy 1925 has two rows of cells in the area between C and RA distal of pterostigma. Vulcagrion Nel &amp; Paicheler 1994 has its veins IR2 and RP3/4 fused basally distal of the subnodus and wing petiolated distal of vein CuP, as for Lithagrion Scudder 1890. Oligoargiolestes Kennedy 1925 diff ers from Electropodagrion in its vein CuP in a more distal position, and...</p></li></ul>