Eteoanisoptera

 Giant dragonfly Petalura gigantea, copyright Leonard Müller.

Belongs within: Anisoptera.
Contains: Gomphoidea, Cavilabiata, Telephlebiidae.

The clade Eteoanisoptera includes the majority of living dragonflies (excluding only the relictual genus Epiophlebia). Members of this group have relatively broad wings that are generally held flat at rest. The larvae are able to move quickly by explosively expelling water from the rectal cavity. Within the Eteoanisoptera, the Petaluroidea are a group of large dragonflies united by a number of forewing synapomorphies including a very narrow postnodal space, with many cells distal from the pterostigma, and a wide subdiscoidal triangle containing a number of cross-veins. The petaluroids are regarded as the sister clade of the remaining dragonflies in the Euanisoptera, but this latter clade is less strongly supported (Nel et al. 2001). Nymphs of modern Petaluridae differ from other dragonflies in burrowing in swampy or marshy soil, emerging from burrows at night to hunt (Richards & Davies 1977).

The Aeshnoidea include the Aeshnidae, hawkers, a group of medium-sized to large, usually strong-flying dragonflies, and their close relatives. Larvae of Aeshnoidea bear sharp lateral spines on at least some abdominal segments, including segment 9 (Watson & O’Farrell 1991). Aeshnids can be distinguished from other living dragonflies by their having wings with vein MA converging on R4 but degenerate distally (Theischinger & Hawking 2006). The Cymatophlebiidae are a family of aeshnoids known from the later Jurassic and early Cretaceous characterised by undulating veins in the wings, particularly vein IR2.

Characters (from Grimaldi & Engel 2005): Wings with pterostigmal brace vein (supporting vein under inner edge of pterostigma); discoidal cell divided into two triangular cells; distinctive anal loop; small, frequently darkened area of veinless membrane present at base of wings near anal region; secondary ‘CuP’ vein associated with expanded anal region in hind wing. Ovipositor vestigial. Naiad able to propel itself using water pressure from rectal chamber.

<==Eteoanisoptera
    |  i. s.: PalaeaeschnaS02
    |         Juragomphus Nel, Bechly et al. 2001 [Juragomphidae]NB01
    |           `--*J. karatauensis Nel, Bechly et al. 2001NB01
    |         Libellulium longialataGE05
    |--Petaluroidea [Petalurida]GE05
    |    |--ProtolindeniidaeRP02
    |    |--Aktassiinae [Aktassiidae]RP02
    |    |    |--Aktassia Pritykina 1968NB01
    |    |    `--Aeschnogomphus Handlirsch 1906NB01
    |    `--PetaluridaeTH06
    |         |--Uropetala carovei (White 1846)HL02
    |         |--Phenes raptor Rambur 1842FN02
    |         `--PetaluraTH06
    |              |--P. gigantea Leach 1815TH06
    |              |--P. hesperia Watson 1958TH06
    |              |--P. ingentissima Tillyard 1908TH06
    |              |--P. litorea Theischinger 1999TH06
    |              `--P. pulcherrima Tillyard 1913TH06
    `--Euanisoptera [Exophytica]RP02
         |--GomphoideaGE05
         `--+--CavilabiataGE05
            `--Aeshnoidea [Aeshnoptera]GE05
                 |  i. s.: Chrysogomphus Ren 1994FB03
                 |           `--*C. beipiaoensis Ren 1994FB03
                 |--EumorbaeschnidaeRP02
                 |--TelephlebiidaeWO91
                 |--MesuropetalaRJ93 [MesuropetalidaeRP02]
                 |    `--M. koehleriRJ93
                 |--CymatophlebiaW13 [CymatophlebiidaeRP02]
                 |    `--C. longialataW13
                 |--Paramesuropetala [Liupanshaniidae]NB01
                 |    `--P. gigantea Bechly et al. 2001NB01
                 `--Aeshnidae [Aeschnidae]TH06
                      |  i. s.: GomphoaeschnaSL89
                      |         *Tetracanthagyna plagiata (Waterhouse 1878) [=Gynacantha plagiata]SL89
                      |         Adversaeschna brevistyla (Rambur 1842)TH06
                      |         BoyeriaA07
                      |         BasiaeshnaA07
                      |         Oligoaeschna Selys Longchamps 1889SL89
                      |           |--*O. modiglianii Selys Longchamps 1889SL89
                      |           `--O. kunigamiensis (Ishida 1972)I92
                      |         Karschia Förster 1900 non Körb. 1865 (ICBN)T08
                      |           |--K. angulata [=Gynacantha angulata]T08
                      |           `--K. cornuta [=Gynacantha cornuta]T08
                      |         Acanthagyna dravidaLM72
                      |--AnactinaeWO91
                      |    |--Hemianax papuensis (Burmeister 1839)WO91, TH06 [=Anax papuensisHL02]
                      |    |--Anaciaeschna Selys 1878WO91, SL89
                      |    |    `--A. jaspidea (Burmeister 1839)TH06 [=Aeschna jaspideaSL89; incl. Anax taitensisSL89]
                      |    `--AnaxTH06
                      |         |--A. amaziliPP72
                      |         |--A. ephippiger [=Hemianax ephippiger]HL02
                      |         |--A. georgius Selys 1872TH06
                      |         |--A. gibbosulus Rambur 1842TH06
                      |         |--A. guttatus (Burmeister 1839)TH06
                      |         |--A. imperatorRD77
                      |         `--A. juniusRD77
                      |--GynacanthaginaeWO91
                      |    |--Agyrtacantha dirupta (Karsch 1889)WO91, TH06
                      |    |--Austrogynacantha Tillyard 1908WO91, T08
                      |    |    `--*A. heterogena Tillyard 1908 [=Gynacantha heterogena de Selys ms]T08
                      |    `--GynacanthaWO91
                      |         |--G. dobsoni Fraser 1951TH06
                      |         |--G. kirbyi Krüger 1898TH06
                      |         |--G. mocsaryi Förster 1898TH06
                      |         |--G. nourlangie Theischinger & Watson 1991TH06
                      |         |--G. rosenbergi Kaup 1867TH06
                      |         `--G. subinterruptaSL89
                      `--AeshninaeWO91
                           |--CaliaeschnaT08
                           `--AeschnaGD00
                                |--A. brevistyla Rambur 1842HL02
                                |--A. canadensisA07
                                |--A. crenataK01
                                |--A. cyaneaGD00
                                |--A. forcipataL02
                                |--A. grandisK01
                                |--A. junceaK01
                                |--A. larvataS02
                                |--A. nigroflavaI92
                                |--A. pryeriK00
                                |--A. sitchensisB-PE07
                                |--A. subarctica Walker 1906I92
                                `--A. thomassoni Kirby 1900K00

*Type species of generic name indicated

References

[A07] Anderson, T. M. 2007. An assessment of water mite parasitism of dragonflies based on museum collections. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 61–69. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[B-PE07] Behan-Pelletier, V. M., & B. Eamer. 2007. Aquatic Oribatida: adaptations, constraints, distribution and ecology. In: Morales-Malacara, J. B., V. M. Behan-Pelletier, E. Ueckermann, T. M. Pérez, E. G. Estrada-Venegas & M. Badii (eds) Acarology XI: Proceedings of the International Congress pp. 71–82. Instituto de Biología and Faculdad de Ciencias, Universidad Nacional Autónoma de México, Sociedad Latinoamericana de Acarología: México.

[FB03] Fleck, G., G. Bechly, X. Martínez-Delclòs, E. Jarzembowski, R. Coram & A. Nel. 2003. Phylogeny and classification of the Stenophlebioptera (Odonata: Epiproctophora). Annales de la Societe Entomologique de France 39 (1): 55–93.

[FN02] Fleck, G., & A. Nel. 2002. The first isophlebioid dragonfly (Odonata: Isophlebioptera: Campterophlebiidae) from the Mesozoic of China. Palaeontology 45 (6): 1123–1136.

[GD00] Giribet, G., D. L. Distel, M. Polz, W. Sterrer & W. C. Wheeler. 2000. Triploblastic relationships with emphasis on the acoelomates and the position of Gnathostomulida, Cycliophora, Plathelminthes, and Chaetognatha: a combined approach of 18S rDNA sequences and morphology. Systematic Biology 49: 539–562.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[HL02] Haacks, M., & A. Lehmann. 2002. Some observations on dragonflies (Insecta: Odonata) throughout New Zealand. Weta 24: 13–17.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[K00] Kirby, W. F. 1900. On a small collection of Odonata (dragonflies) from Hainan, collected by the late John Whitehead. Annals and Magazine of Natural History, series 7, 5: 530–539, pl. 12.

[K01] Klapálek, F. 1901. Neuropteroidák [Neuropteroiden]. In: Horváth, G. (ed.) Zichy Jenő Gróf Harmadik Ázsiai Utazása [Dritte Asiatische Forschungsreise des Grafen Eugen Zichy] vol. 2. Zichy Jenő Gróf Harmadik Ázsiai Utazásának Állattani Eredményei [Zoologische Ergebnisse der Dritten Asiatischen Forschungsreise des Grafen Eugen Zichy] pp. 203–221. Victor Hornyánszky: Budapest, and Karl W. Hierseman: Leipzig.

[LM72] Lahiri, A. R., & T. R. Mitra. 1972. A note on Acanthagyna dravida (Lieftinck) [Insecta: Odonata: Aeshnidae]. Journal of the Bombay Natural History Society 69 (2): 438–439.

[L02] Latreille, P. A. 1802. Histoire Naturelle, générale et particulière des crustacés et des insectes vol. 3. Familles naturelles des genres. F. Dufart: Paris.

[NB01] Nel, A., G. Bechly, X. Martínez-Delclòs & G. Fleck. 2001. A new family of Anisoptera from the Upper Jurassic of Karatau in Kazakhstan (Insecta: Odonata: Juragomphidae n. fam.) Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 314: 1–9.

[PP72] Parkin, P., D. T. Parkin, A. W. Ewing & H. A. Ford. 1972. A report on the arthropods collected by the Edinburgh University Galapagos Islands expedition, 1968. Pan-Pacific Entomologist 48 (2): 100–107.

[RP02] Rasnitsyn, A. P., & L. N. Pritykina. 2002. Superorder Libellulidea Laicharting, 1781. Order Odonata Fabricius, 1792. The dragonflies. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 97–104. Kluwer Academic Publishers: Dordrecht.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[SL89] Selys Longchamps, E. de. 1889. Odonates de Sumatra comprenant les espèces recueillies à Pulo Nias par M. le D.r E. Modigliani. Annali del Museo Civico di Storia Naturale di Genova, Serie 2a, 7: 444–484.

[S02] Sinitshenkova, N. D. 2002. Ecological history of the aquatic insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 388–426. Kluwer Academic Publishers: Dordrecht.

[TH06] Theischinger, G., & J. Hawking. 2006. The Complete Field Guide to Dragonflies of Australia. CSIRO Publishing: Collingwood (Victoria).

[T08] Tillyard, R. J. 1908. On the new genus Austrogynacantha [Neuroptera: Odonata] with description of species. Proceedings of the Linnean Society of New South Wales 33: 423–431, pl. 5.

[WO91] Watson, J. A. L., & A. F. O’Farrell. 1991. Odonata (dragonflies and damselflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers 2nd ed. vol. 1 pp. 294–310. Melbourne University Press: Carlton (Victoria).

[W13] Witton, M. P. 2013. Pterosaurs: Natural History, Evolution, Anatomy. Princeton University Press: Princeton (New Jersey).

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