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Paleontologia Lombarda PALEONTOLOGIA LOMBARDA Nuova serie Volume X Andre Nel, Giinter Bechly, Edmund Jarzembowski & Xavier Martinez-Delclos A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida) Societa Italiana di Scienze Naturali Museo Civico di Storia Naturale di Milano Milano 1998 SerieI Seriem STOPPANI A., 1860-1865 - Geologie et Paleontologie des cou­ STOPPANI A., 1858-1860 - Les Petriffcations d'Esino ou ches l'l Avicula contorta en Lombardie. 266 pp., 59 tavv. (out of description des fossiles appartenant au depot triasique supe­ print) rieur des environs d'Esino en Lombardie.152 pp.,31 tavv. (out of print) SerieIV MENEGHINI G., 1867-1881 - Monographie des fossiles du cal­ SerieD caire rouge ammonitique (Lias Superieur) de Lombardie et de I'Apennin Central. 242 pp., 31 tavv. (out of print) CoRNALIA E., 1858-1871 - Mammiferes fossiles de Lombardie. MENEGHINI G., 1867-1881 - Fossiles du Medolo. 56 pp., 7 tavv. 95 pp., 28 tavv. (out of print) (out of print) NUOVASERIE Volume I Volume VI GARASSINO A. & TERUZZI G., 1993 - A new decapod NOSOTII S. & PINNA G., 1996 - Osteology of the skull crustacean assemblage from the Upper Triassic of of Cyamodus kuhnschnyderi Nosotti & Pinna 1993 Lombardy (N. Italy). 27 pp. , 5 tab. (Reptilia, Placodontia). 42 pp. VolumeD Volume VII MAZIN J.M. & PINNA G. eds. , 1993 - Evolution, eco­ DAL SASSO C. & PINNA G., 1997- Aphanizocnemus logy and biogeography of the Triassic Reptiles. 170 libanensis n. gen. n. sp. , a new dolichosaur (Reptilia, pp. Varanoidea) from the Upper Cretaceous of Lebanon. 31 pp. Volume Ill Volume VIII GARASSINO A., 1994- The macruran decapod cru­ staceans of the Upper Cretaceous of Lebanon. 27 HARTENBERGER J-L. , 1998 - The Cenozoic radiation pp. , XIII pi. of Mammals: some comments on the shape and tempo of a major evolutionary event. 21 pp. Volume IV Volume IX DAL SASSO C. & PINNA G., 1996 - Besanosaurus lep­ torhynchus n. gen. n. sp. , a new shastasaurid ichthyo­ DUFFIN C. , 1997- Ostenose/ache stenosoma n.g. n.sp. , saur from the Middle Triassic of Besano a new neoselachian shark from the Sinemurian (Lombardy, N. Italy). 22 pp. (Early Jurassic) of Osteno (Lombardy, Italy). 27 pp., XII pi. Volume V GALL J-C ed. , 1996 - Triassic insects of Western Europe. 60 pp. Andre Nel (*), Giinter Bechly (**), Edmund Jarzembowski (***) & Xavier Martinez-Delclos (****) (*) Museum National d'Histoire Naturelle, Paris (**) Eberarhard-Karls-UniversiUit, Tiibingen (***) Maidstone Museum & Art Gallery, Kent (****) Universitat de Barcelona A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida) Nuova serie - Volume X Societa Italiana di Scienze Naturali Museo Civico di Storia Naturale di Milano Milano 1998 CONTENTS Introduction ................................................................................ Pag. 4 The alleged plesiomorphy Systematics and taxonomy ......................................... Pag. 6 of semi-terrestrial larvae ............................................... Pag. 59 Cladistic analysis of petalurida .............................. Pag. 48 Fossil larvae of Petalurida ........................................... Pag. 60 Clad istic analysis ................................................................... Pag. 48 Scarcity in the fossil record of Petalurida ..... Pag. 60 Character listing ................................................................... Pag. 50 Stem-group representatives of extant Ty pe specimens ....................................................................... Pag. 54 Petaluridae ................................................................................ Pag. 60 Phylogeny and evolution of petalurida............ Pag. 55 Proposed phylogenetic system of Petalurida Pag. 60 Systematic position of the Mesuropetalidae Pag. 55 Anisoptera incertae sedis ........................................... Pag. 64 Systematic position of the Cymatophlebiidae. Pag. 55 Anisoptera incertae sedis (probably no Phylogenetic position of Petalurida within Petalurida) ................................................................................ Pag. 64 Anisoptera .................................................................................. Pag. 56 Historical biogeography ............................................... Pag. 65 Plesiomorphic ground-plan characters of Acknowledgements ............................................................ Pag. 65 Petalurida ................................................................................... Pag. 58 Literature ..................... .............................................................. Pag. 66 Editorial board Jan Bergstrom (Sweden), Derek Briggs (UK), Jean Claude Gall (France), Jean-Louis Hartenberger (France), Peter Wellnhofer (Germany) © Societa ltaliana di Scienze Naturali e Museo Civico di Storia Naturale di Milano Corso Venezia, 55- 20121 Milano Registrato al Tribunale di Mllano al n. 137 del 22/2/1992 Direttore responsabile Giovanni Pinna Grafica editoriale Michela Mura Stampa Litografia Solari, Peschiera Borromeo, luglio 1998 ISSN 1121-3361 Andre Nel, Giinter Bechly, Edmund Jarzembowski & Xavier Martinez-Del cl os A revision of the fossil petalurid dragonflies (Insecta: Odonata: Anisoptera: Petalurida) Abstract - A new family, genus and species of Petalurida, Cretapeta/ura brasiliensis gen. nov. et sp. nov. (Cretapetaluri­ dae fam. nov.) is described from the Lower Cretaceous Santana Formation of the Araripe Basin of Brazil, and a new sub­ family, genus and species of Petalurida, Pseudocymatophlebia henn.igi gen. nov. et sp. nov. (Pseudocymatophlebiinae subfam. nov. in Aktassiidae) is described from the Lower Cretaceous Weald Clay of England. A new specie Aktassia pritykinae sp. nov. is described from the Lower Cretaceous of Mongolia. 1l1e description of new material enables us to revise the phyla­ genetic position of the genera Protolindenia Deichmliller 1886, Aeschnogomphus Handlirsch 1906, Mesuropetala Handlirsch 1906, and Cymatophlebia Deichmliller 1886 from the Upper Jurassic of Germany, and to designate neotypes for Protolinde­ nia willei and Mesuropetala koehleri. Aeschnogomphus and Aktassia Pritykina 1968 are considered to be sister-genera with­ in the Petalurida - Aktassiidae (subfamily Aktassiinae stat. nov.). Aeschnogomphus buchi (Hagen 1848) is recognized as valid name for Aeschnogomphus charpentieri (Hagen 1848). Mesuropetala, formerly considered to be a petalurid, is regard­ ed as a basal Aeshnoptera; and Protolindenia, formerly considered to be a gomphid, is transferred to the Petalurida, as most basal member of the stem-group of Petaluridae. The phylogenetic positions of Mesuropeta/a auliensis Pritykina 1968, Mesuropetala costalis Pritykina 1968, Protolin.denia aktassica Pritykina 1968 (in Kazakhophlebiella gen. nov. et comb. nov.) and Proto/indenia deichmuel/eri Pritykina 1968 (in Pritykiniella gen. nov. et comb. nov.) (all Upper Jurassic taxa from Karatau, Turkestan,Russian Federation), are discussed. Also, the phylogenetic positions of Miopetalura shanwangica Zhang 1989 and Miopetalura orientalis (Hong 1985) (Middle Miocene of China) are discussed and these taxa are transferred from the Petaluridae to the Gomphides- Lindeniinae and Anisoptera incertae sedis respectively. The English Lower Cretaceous Aeschnopsis perampla (Brodie 1845) and Cy rnatophlebiopsis pseudobubas Handlirsch 1939 are revised, synonymised and considered to belong to Anisoptera incertae seclis. The Lower Cretaceous genus Necrogornphus Campion 1923 with two species N. petrificatus (Hagen 1850) and N. jurassicus (Giebel 1850) is revised and also referred to Anisoptera incertae sedis. l11e phylogenetic positions of Protolindenia, Aeschnogomphus, Aktassia, Pseudocymatophlebia gen. nov., and Cretapetalura gen. nov. within the Petalurida are discussed and a phylogenetic analysis of the fossil and extant Petalurida is presented. The Petalurida are identified as sister-group of all remaining extant Anisoptera (Euanisoptera). The new phylogenetic system of Anisoptera by Bechly (1996) is confirmed, and new phylogenetic definitions of the taxon names of Petalurida are proposed. 1l1e evolution and historical biogeography of Petalurida is discussed. Resume- Revision des libellules petalurides fossiles. (Insecta: Odonata: Anisoptera: Petalurida). Une nouvelle famille, genre et espece de Petalurida Cretapetalura brasiliensis gen. nov. et sp. nov., est decrit du Cretace inferieur de la Formation Santana du bassin lacustre d'Araripe en Bresil, et une nouvelle sous-famille, genre et espece de Petalurida, Pseudocymatophlebia hennigi gen. nov. et sp. nov. (Pseudocymatophlebiinae subfam. nov. in Aktassiidae) est de­ crit du Cretace inferieur du 'Weald Clay' en Angleterre. Une nouvelle espece Aktassia pritykinae sp. nov. est decrite du Cre­ tace inferieur de Mongolie. De plus, la description de nouveaux specimens nous permet de reviser la position phylogene­ tique des genres Prowlindenia Deichmliller 1886, Aeschnogomphus Handlirsch 1906, Mesuropeta/a Handlirsch 1906 et Cy­ matophlebia Deichmliller 1886 du Jurassique superieur d'All emagne, et de designer des neotypes pour Protolirulenia willei et Mesuropeta/a koehleri. Aeschnogomphus et Aktassia Pritykina 1968 sont consideres comme des genres freres dans les Pe­ talurida - Aktassiidae. Aeschnogomphus buchi (Hagen 1848) est reconnu comme nom valide en remplacement d'Aeschno­ gornphus charpentieri (Hagen 1848). Mesuropeta/a, anterieurement range dans les Petalurida, est considere comme un Ae­ shnoptera. Protolindenia, anterieurement
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