A revision of the Upper Jurassic-Lower Cretaceous dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Insecta, Odonatoptera, Panodonata) Günther FLECK Département Histoire de la Terre, USM 0203 and CNRS-Muséum UMR 5143, Muséum national d’Histoire naturelle, 45 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Günter BECHLY Staatliches Museum für Naturkunde Stuttgart, Rosenstein 1, D-70191 Stuttgart (Germany) [email protected] Xavier MARTÍNEZ-DELCLÒS Departament d’Estratigrafia, Paleontologia i Geociències Marines, Facultat de Geologia, Universitat de Barcelona, E-08071 Barcelona (Spain) [email protected] Edmund A. JARZEMBOWSKI Maidstone Museum and Bentlif Art Gallery, St Faith’s St, Maidstone, Kent ME14 1LH (UK) and Postgraduate Research Institute for Sedimentology & PRIS, University of Reading, Reading RG6 2AB (UK) [email protected] André NEL Département Histoire de la Terre, USM 0203 and CNRS-Muséum UMR 5143, Muséum national d’Histoire naturelle, 45 rue Buffon, F-75231 Paris cedex 05 (France) [email protected] Fleck G., Bechly G., Martínez-Delclòs X., Jarzembowski E. A. & Nel A. 2004. — A revision of the Upper Jurassic-Lower Cretaceous dragonfly family Tarsophlebiidae, with a discussion on the phylogenetic positions of the Tarsophlebiidae and Sieblosiidae (Insecta, Odonatoptera, Panodonata). Geodiversitas 26 (1) : 33-60. GEODIVERSITAS • 2004 • 26 (1) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 33 Fleck G. et al. ABSTRACT The Upper Jurassic-Lower Cretaceous dragonfly family Tarsophlebiidae is revised. The type species of the type genus Tarsophlebia Hagen, 1866, T. eximia (Hagen, 1862) from the Upper Jurassic Solnhofen Limestones, is redescribed, including important new information on its head, legs, wings, anal appendages and male secondary genital apparatus. The type specimen of Tarsophlebiopsis mayi Tillyard, 1923 is regarded as an aberrant or unusually preserved Tarsophlebia eximia. One new species of Tarsophlebia and three new species of Turanophlebia are described, i.e. Tarsophlebia minor n. sp., Turanophlebia anglicana n. sp., T. mongolica n. sp., and T. vitimensis n. sp. A new combination is proposed for Turanophlebia neckini (Martynov, 1927) n. KEY WORDS comb. The phylogenetic relationships of the Mesozoic Tarsophlebiidae are Insecta, Odonatoptera, discussed on the basis of new body and wing venation characters. The present Zygoptera, analysis supports a rather derived position for the Tarsophlebiidae, as sister Epiproctophora, group of the the Epiproctophora rather than of (Zygoptera + Epiproc- Tarsophlebiidae, Sieblosiidae, tophora). Also, through the present discussion, the Oligo-Miocene family Upper Jurassic-Lower Sieblosiidae seems to be more closely related to the Epiproctophora than to Cretaceous, Cenozoic, the Zygoptera. But the present study and previous analyses suffer of the lack fossil, of informations concerning the more inclusive groups of Odonatoptera, viz. copulation, Protozygoptera, Triadophlebiomorpha, Protanisoptera, etc. The significance phylogeny, revision, of the tarsophlebiid secondary male genital apparatus for the reconstruction new species. of the evolution of odonate copulation is discussed. RÉSUMÉ Révision des Tarsophlebiidae du Jurassique supérieur et Crétacé inférieur, discus- sion sur les positions phylogénétiques des Tarsophlebiidae et des Sieblosiidae (Insecta, Odonatoptera, Panodonata). La famille de libellules mésozoïques des Tarsophlebiidae est révisée. L’espèce type du genre type Tarsophlebia Hagen, 1866, T. eximia (Hagen, 1862) du cal- caire lithographique du Jurassique supérieur de la Bavière, est redécrite, in- cluant de nouvelles données sur les structures de la tête, des pattes, des ailes, des appendices anaux et des pièces génitales secondaires du mâle. Le type de Tarsophlebiopsis mayi Tillyard, 1923 est considéré comme un spécimen aberrant ou curieusement préservé de Tarsophlebia eximia. Une nouvelle espèce de Tarsophlebia et trois nouvelles espèces de Turanophlebia sont décrites : Tarsophlebia minor n. sp., Turanophlebia anglicana n. sp., T. mongolica n. sp. et T. vitimensis n. sp. Une nouvelle combinaison est proposée pour Turanophlebia MOTS CLÉS neckini (Martynov, 1927) n. comb. Les relations phylogénétiques des Insecta, Tarsophlebiidae sont discutées sur la base des nouveaux caractères du corps et Odonatoptera, Zygoptera, des ailes. Notre analyse conclut à une probable position des Tarsophlebiidae Epiproctophora, comme groupe frère des Epiproctophora, plutôt que des (Zygoptera + Tarsophlebiidae, Sieblosiidae, Epiproctophora). Au travers de la présente discussion, il ressort que la famille Jurassique supérieur- cénozoïque des Sieblosiidae semble plus proche des Epiproctophora que des Crétacé inférieur, Zygoptera. Il apparaît aussi clairement que toutes ces analyses phylogénétiques Cénozoïque, fossile, souffrent du manque de données concernant les groupes plus basaux, comme copulation, les Protozygoptera, Triadophlebiomorpha ou Protanisoptera. L’intérêt des phylogénie, pièces génitales secondaires des mâles des Tarsophlebiidae pour l’analyse de révision, nouvelles espèces. l’évolution de la copulation chez les Odonatoptera est discuté. 34 GEODIVERSITAS • 2004 • 26 (1) Revision of Tarsophlebiidae (Odonatoptera) INTRODUCTION Bq series of cross-veins between RP1+2, IR2 and oblique vein; The Tarsophlebiidae Handlirsch, 1906 is one of C costal vein or costa; Cr nodal cross-vein; the most enigmatic Mesozoic family of Odona- CuA Cubitus Anterior; toptera. Its phylogenetic relationships with the Zy- CuAb first posterior branch of Cubitus goptera and the Epiproctophora remain very un- Anterior; certain. Nel et al. (1993) considered them as the CuP Cubitus Posterior; most inclusive lineage within the “Anisozygoptera” IR intercalary vein of radial area; MA Median Anterior; + Anisoptera (= Epiproctophora). Bechly (1996) MAb first posterior branch of Median and Rehn (2003) proposed to place them as sister Anterior; group of the (Zygoptera + Epiproctophora). After MP Median Posterior; our recent studies at PIN and MCZ, we could re- RA Radius Anterior; vise and describe several species. This study also re- RP Radius Posterior; vealed that the alleged calopterygoid-like anal ap- ScA Subcosta Anterior ScP Subcosta Posterior; pendages of Tarsophlebia, with apparently two pairs of claspers, are clearly based on misinterpreta- tions due to artifacts of preservation. Indeed Tarso- phlebia definitely does not possess zygopteroid but MATERIAL AND METHODS rather unique appendages. There are no visible paraprocts, the epiproct must have been reduced All drawings were made with a camera lucida and or inconspicuous since it is not visible in any speci- a binocular microscope. The nomenclature of the men, and the cerci are very long, with a double- dragonfly wing venation is based on the interpre- barrelled basal petiole and a curious distal plate- tations of Riek & Kukalová-Peck (1984), amen- like expansion. The broken double-barrelled ded by Nel et al. (1993) and Bechly (1996). The petioles of the two cerci have been commonly mi- systematic analysis is based on the principles of sinterpreted as two pairs of claspers, while the dis- consequent phylogenetic systematics (sensu tal plates have been overlooked or regarded as pre- Hennig 1966, 1969, 1981), especially character servational artifacts. This new result is of particular polarisation based on outgroup comparison. phylogenetic relevance, since the previous inter- pretation of the appendages would have represen- ted a putative synapomorphy with Zygoptera. We SYSTEMATICS comment on the characters that are used for the phylogenetic position of the Tarsophlebiidae. Order ODONATOPTERA Martynov, 1932 Suborder PANODONATA Bechly, 1996 ABBREVIATIONS Institutions Family TARSOPHLEBIIDAE Handlirsch, 1906 BSPGM Bayerische Staatssamlung für Paläon- tologie und Historische Geologie, Tarsophlebiidae Handlirsch, 1906: 467, 468, 580, Munich; 581. JME Jura-Museum, Eichstätt; MCZ Museum of Comparative Zoology, TYPE GENUS.— Tarsophlebia Hagen, 1866, original Harvard University, Cambridge; designation of Handlirsch (1906) by monotypy. PIN Arthropod Laboratory, Palaeonto- INCLUDED GENERA.— Tarsophlebia Hagen, 1866 (= logical Institute, Academy of Science Tarsophlebiopsis Tillyard, 1923) and Turanophlebia of Russia, Moscow; Pritykina, 1968. Since Bechly (1996, 1997) demon- Names of body and wing structures strated that the genera Euthemis Pritykina, 1968 (Upper Jurassic) and Sphenophlebia Bode, 1953 AA Analis Anterior; (Upper Liassic) are isophlebioid “anisozygoptères” and Ax1 and Ax2 primary antenodal cross-veins; thus not related to Tarsophlebiidae, there is no more GEODIVERSITAS • 2004 • 26 (1) 35 Fleck G. et al. evidence for a Lower Jurassic occurrence of Liassic of Gloucestershire in England was correctly Tarsophlebiidae and also no need for a further redun- transferred to the heterophlebioid genus Liassophlebia dant taxon Tarsophlebioidea or Tarsophlebioptera. by Tillyard (1925). STRATIGRAPHIC AND GEOGRAPHIC RANGE. — Only EMENDED DIAGNOSIS. — Nel et al. (1993) proposed a known from the Upper Jurassic and Lower Cretaceous diagnosis, erroneous in some points. We emend it as of Eurasia. follows: primary antenodal braces stronger than sec- ondaries, as in other Tarsophlebiidae (contra Nel et al. EMENDED DIAGNOSIS. — Nel et al. (1993) proposed a 1993); cubito-anal areas of fore and hind wings with diagnosis