Cavilabiata

Alpine redspot dragonfly Austropetalia tonyana, copyright Gunther Theischinger.

Belongs within: Eteoanisoptera.
Contains: Brachystigmata.

The Cavilabiata are a clade of dragonflies uniting the Cordulegastridae, Libellulidae and related families. Other than the basal Cordulegastridae, most members of this clade belong to the subclade Cristotibiata, united by a rather stout pterostigma that is not parallel-sided, a shortened hindwing CuA, and an enlarged anal loop (Bechly & Ueda 2002). Species of the families Neopetaliidae, Archipetalia and Austropetalia have the front part of each pair of wings marked with a series of rounded, reddish brown spots.

Synapomorphies (from Bechly & Ueda 2002): Discoidal triangles at least somewhat longitudinal elongate in both wings, especially in the hind wing; distal part of antesubnodal area free of crossveins (“cordulegastrid gap”); “gaff” (basal CuA before its branching) of hindwing at least slightly prolonged; RP3/4 and MA slightly undulating in both pairs of wings.

<==Cavilabiata
    |--Cordulegaster I92 [Cordulegastrida BU02, Cordulegastridae, Cordulegastroidea]
    |    `--C. boltoni I92
    `--Cristotibiata BU02
         |--Brachystigmata BU02
         `--+--Phyllopetalia WO91 [Neopetaliidae BU02]
            |--Archipetalia WO91 [Archipetaliidae TH06]
            |    `--A. auriculata Tillyard 1917 TH06
            `--Austropetalia WO91 [Austropetaliidae TH06]
                 |--A. patricia (Tillyard 1910) TH06
                 `--A. tonyana Theischinger 1995 TH06

*Type species of generic name indicated

REFERENCES

[BU02] Bechly, G., & K. Ueda. 2002. The first fossil record and first New World record for the dragonfly clade Chlorogomphida (Insecta: Odonata: Anisoptera: Araripechlorogomphidae n. fam.) from the Crato Limestone (Lower Cretaceous, Brazil). Stuttgarter Beiträge zur Naturkunde Serie B (Geologie und Paläontologie) 328: 1–11.

[I92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

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

[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 2^nd ed. vol. 1 pp. 294–310. Melbourne University Press: Carlton (Victoria).

Last updated: 29 May 2022.

Telephlebiidae

Male alpine darner Austroaeschna flavomaculata, copyright Toby Hudson.

Belongs within: Anisoptera.

The Telephlebiidae, southern darners, are a group of medium-sized to very large dragonflies found only in eastern and south-western Australia (Theischinger & Hawking 2006).

Characters (from Theischinger & Hawking 2006): Anterior portion of wing mostly clear except pterostigma, rarely marked with brown band(s) or rounded nodal spot; vein Rsp1 only slightly curved, almost parallel with vein IR3; vein MA unbroken distally, runs parallel to or diverges from R4. Larvae profoundly elongate; prementum flat, lacks premental setae; mid-dorsal abdominal spines usually absent, rarely present; some abdominal segments, including segment 9, armed with lateral spines; epiproct strongly tapered throughout with apex commonly pointed.

<==Telephlebiidae TH06
    |--Acanthaeschna victoria Martin 1901 TH06
    |--Antipodophlebia asthenes (Tillyard 1916) TH06
    |--Dendroaeschna conspersa (Tillyard 1907) TH06
    |--Dromaeschna forcipata ZS10
    |--Austrophlebia TH06
    |    |--A. costalis (Tillyard 1907) TH06
    |    `--A. subcostalis Theischinger 1996 TH06
    |--Spinaeschna TH06
    |    |--S. tripunctata (Martin 1901) TH06
    |    `--S. watsoni Theischinger 1982 TH06
    |--Notoaeschna TH06
    |    |--N. geminata Theischinger 1982 TH06
    |    `--N. sagittata (Martin 1901) TH06
    |--Telephlebia TH06
    |    |--T. brevicauda Tillyard 1916 TH06
    |    |--T. cyclops Tillyard 1916 TH06
    |    |--T. godeffroyi Selys 1883 TH06
    |    |--T. tillyardi Campion 1916 TH06
    |    |--T. tryoni Tillyard 1917 TH06
    |    `--T. undia Theischinger 1985 TH06
    `--Austroaeschna TH06
         |--A. anacantha Tillyard 1908 TH06
         |--A. atrata Martin 1909 TH06
         |--A. christine Theischinger 1993 TH06
         |--A. cooloola Theischinger 1991 TH06
         |--A. eungella Theischinger 1993 TH06
         |--A. flavomaculata Tillyard 1916 TH06
         |--A. forcipata (Tillyard 1907) TH06
         |--A. hardyi Tillyard 1917 TH06
         |--A. inermis Martin 1901 TH06
         |--A. muelleri Theischinger 1982 TH06
         |--A. multipunctata (Martin 1901) TH06
         |--A. obscura Theischinger 1982 TH06
         |--A. parvistigma (Selys 1883) TH06
         |--A. pulchra Tillyard 1909 TH06
         |--A. sigma Theischinger 1982 TH06
         |--A. speciosa Sjöstedt 1917 TH06
         |--A. subapicalis Theischinger 1982 TH06
         |--A. tasmanica Tillyard 1916 TH06
         |--A. unicornis (Martin 1901) TH06
         |    |--A. u. unicornis TH06
         |    `--A. u. pinheyi TH06
         `--A. weiskei (Förster 1908) TH06

*Type species of generic name indicated

REFERENCES

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

[ZS10] Zborowski, P. & R. Storey. 2010. A Field Guide to Insects in Australia. Reed New Holland: Sydney.

Anisoptera

Part and counterpart of Aeschnidium heishankowense, from here.

Belongs within: Euepiproctophora.
Contains: Eteoanisoptera.

The Anisoptera as recognised here includes the modern dragonflies (Eteoanisoptera) and a number of close fossil relatives, united by the possession of non-petiolate wings. Excluding the basal Henrotayia marci, whose affinities are somewhat uncertain, Anisoptera are also supported by fore wings with the nodal Cr nearly perpendicular to RA and ScP, and with a pseudo-IR1 vein; the presence of an 'anisopteroid vein' (AA + CuA)b in the hind wing; and a transverse anal triangle in the male. The Aeschnidiidae are united with modern dragonflies by the long hypertriangle in the fore wing in contrast to the short hypertriangle in Liassogomphidae (Fleck et al. 2003).

<==Anisoptera [Pananisoptera]
    |  i. s.: Sona R02 [Sonidae FB03]
    |           `--S. nectes Pritykina 1986 R02
    |         Petaliaeschna fletcheri Fraser 1927 FB03
    |--Henrotayia Fleck, Bechly et al. 2003 [Henrotayiidae] FB03
    |    `--*H. marci Fleck, Bechly et al. 2003 FB03
    `--+--Liassogomphidae [Gomphitidae] FB03
       |    |--Liassogomphus brodiei (Buckman 1843) FB03
       |    |--Elattogomphus latus Bode 1953 NB01
       |    |--Proinogomphus bodei (Handlirsch 1939) NB01
       |    `--Phthitogomphus angulatus (Handlirsch 1939) NB01
       `--Neoanisoptera NB01
            |--Eteoanisoptera FB03
            `--Aeschnidiidae FB03
                 |--Aeschnidium heishankowense ZBH03
                 |--Huizhougenia Lin 1980 KS02
                 |--Nothomacromia [Nothomacromiidae] S02
                 |--Aeschnidopsis flindersiensis RJ93
                 `--Hebeiaeschnidia fengningensis RJ93

*Type species of generic name indicated

REFERENCES

[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.

[KS02] Kluge, N. Yu., & N. D. Sinitshenkova. 2002. Order Ephemerida Latreille, 1810. The true mayflies (=Ephemeroptera Hyatt et Arms, 1891 (s. l.); =Euephemeroptera Kluge, 2000. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 89–97. Kluwer Academic Publishers: Dordrecht.

[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.

[R02] Rasnitsyn, A. P. 2002. Special features of the study of fossil insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 8–12. Kluwer Academic Publishers: Dordrecht.

[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.

[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.

[ZBH03] Zhou, Z., P. M. Barrett & J. Hilton. 2003. An exceptionally preserved Lower Cretaceous ecosystem. Nature 421: 807–814.

Last updated: 29 May 2022.

Hemicorduliidae

Australian emerald Hemicordulia australiae, copyright Fir0002/Flagstaffotos.

Belongs within: Eteoanisoptera.

The Hemicorduliidae, round-winged emeralds, are a group of small to medium-sized dragonflies most commonly black and yellow in coloration (Theischinger & Hawking 2006).

Characters (from Theischinger & Hawking 2006): Forewing with hypertriangles generally free; bridge crossvein aligned with subnodus. Males with tibial keels. Larvae with ladle-shaped labium with premental and palpal setae, well-developed palpal dentations bearing groups of setae; prementum with distinct ventro-basal midline or groove; legs long; abdominal segment 10 somewhat sunken into segment 9.

<==Hemicorduliidae
    |--Procordulia TH06
    |    |--P. affinis (Selys 1871) TH06
    |    |--P. grayi (Selys 1871) HL02
    |    |--P. jacksoniensis (Rambur 1842) TH06
    |    `--P. smithii (White 1846) HL02
    `--Hemicordulia TH06
         |--H. australiae (Rambur 1842) TH06
         |--H. continentalis Martin 1907 TH06
         |--H. flava Theischinger & Watson 1991 TH06
         |--H. intermedia (Selys 1871) TH06
         |--H. kalliste Theischinger & Watson 1991 TH06
         |--H. koomina Watson 1969 TH06
         |--H. mindana I92
         |    |--H. m. mindana I92
         |    `--H. m. nipponica Asahina 1980 I92
         |--H. novaehollandiae (Selys 1871) TH06
         |--H. ogasawarensis Oguma 1913 I92
         |--H. okinawensis Asahina 1947 I92
         |--H. superba Tillyard 1911 TH06
         `--H. tau (Selys 1871) TH06

*Type species of generic name indicated

REFERENCES

[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.

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

Euodonata

Counterpart, part and interpretive diagram of holotype of Saxonagrion minutus, from Nel et al. (1999).

Belongs within: Pandiscoidalia.
Contains: Zygoptera, Euepiproctophora, Isophlebioptera.

The Euodonata is the crown group of dragonflies and damselflies. Synapomorphies of this clade include the formation of the posterior arculus at the separation of veins RP and MA, and the presence of the subdiscoidal vein (Grimaldi & Engel 2005). The earliest record of this clade is from the Late Permian, with the stem-damselfly Saxonagrion minutus being found in the Salagou Formation of France (Nel et al. 1999).

<==Euodonata
    |--+--Zygoptera FB03
    |  `--Saxonagrion NB01 [Saxonagrionidae GE05]
    |       `--S. minutus Nel et al. 1999 NB01
    `--Epiprocta [Epiproctophora] GE05
         |  i. s.: Oreopteridae GE05
         |         Turanothemistidae GE05
         |--Euepiproctophora FB03
         `--+--Steleopteridae RP02
            `--Isophlebioptera FB03

Euodonata incertae sedis:
  Shurabiola Pritykina 1980 FN02
  Mortonagrion hirosei Asahina 1972 I92
  Rhipidolestes I92
    |--R. aculeata I92
    |    |--R. a. aculeata I92
    |    |--R. a. hiraoi I92
    |    `--R. a. yakusimensis I92
    `--R. okinawanus Asahina 1951 I92
  Boninthemis insularis (Matsumura 1913) I92
  Lyriothemis I92
    |--L. elegantissima I92
    `--L. tricolor Ris 1916 I92
  Bayadera brevicauda I92
    |--B. b. brevicauda I92
    `--B. b. ishigakiana Asahina 1964 I92
  Matrona basilaris I92
    |--M. b. basilaris I92
    |--M. b. japonica Foerster 1897 I92
    `--M. b. nigripectus I92
  Asiagomphus I92
    |--A. amamiensis (Asahina 1962) I92
    |    |--A. a. amamiensis I92
    |    `--A. a. okinawanus (Asahina 1964) I92
    `--A. yayeyamensis (Oguma 1926) I92
  Leptogomphus I92
    |--L. sauteri I92
    |    |--L. s. sauteri I92
    |    `--L. s. formosanus I92
    `--L. yayeyamensis Oguma 1926 I92
  Macromidia ishidai Asahina 1964 I92
  Cercion calamorum I92
  Boninagrion ezoin Asahina 1952 I92
  Stylogomphus I92
    |--S. ryukyuanus I92
    |    |--S. r. ryukyuanus I92
    |    `--S. r. asatoi Asahina 1972 I92
    `--S. shirozui I92
         |--S. s. shirozui I92
         `--S. s. watanabei Asahina 1984 I92
  Planaeschna I92
    |--P. ishigakiana Asahina 1951 I92
    |    |--P. i. ishigakiana I92
    |    `--P. i. nagaminei Asahina 1988 I92
    `--P. risi I92
         |--P. r. risi I92
         `--P. r. sakishimana Asahina 1964 I92
  Trapezostigma loewii TDC06
  Argiolestes minimus ASD05

*Type species of generic name indicated

REFERENCES

[ASD05] Armstrong, K. N., A. W. Storey & P. M. Davies. 2005. Effects of catchment clearing and sedimentation on macroinvertebrate communities of cobble habitat in freshwater streams of southwestern Australia. Journal of the Royal Society of Western Australia 88 (1): 1-11.

[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.

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

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

Nel, A., G. Gand, G. Fleck, O. Bethoux, J. Lapeyrie & J. Garric. 1999. Saxonagrion minutus nov. gen. et sp., the oldest Damselfly from the Upper Permian of France (Odonatoptera, Panodonata, Saxonagrionidae nov. fam.) Geobios 32 (6): 883-888.

[NB01] Nel, A., O. Bethoux, G. Bechly, X. Martínez-Delclòs & F. Papier. 2001. The Permo-Triassic Odonatoptera of the “protodonate” grade (Insecta: Odonatoptera). Annales de la Société Entomologique de France (N.S.) 37 (4): 501-525.

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

[TDC06] Timms, B. V., B. Datson & M. Coleman. 2006. The wetlands of the Lake Carey catchment, northeast Goldfields of Western Australia, with special reference to large branchiopods. Journal of the Royal Society of Western Australia 89 (4): 175-183.

Pandiscoidalia

Holotype of Reisia gelasii, from Museum Terra Triassica.

Belongs within: Odonatoptera.
Contains: Triadophlebiomorpha, Euodonata.

The Pandiscoidalia is a clade of odonatopterans including the living damselflies and dragonflies, as well as a number of closely related stem taxa.

Synapomorphies (from Nel et al. 2001): Wings with highly developed nodus with kink in ScP; nodal furrow present; nodal and subnodal crossveins aligned; primary antenodal crossvein Ax1 present.

<==Pandiscoidalia [Triadophlebioptera]
    |--Triadotypidae [Reisiidae, Triadotypomorpha] NB01
    |    |--Triadotypus Grauvogel & Laurentiaux 1952 NB01
    |    |    |--*T. guillaumei Grauvogel & Laurentiaux 1952 [=Reisia guillaumei] NB01
    |    |    `--T. nana Bechly 1997 [=T. guillaumei var. nana] NB01
    |    `--Reisia Handlirsch 1912 NB01
    |         |--*R. gelasii (Reis 1909) [=Handlirschia gelasii] NB01
    |         `--R. sodgianus (Pritykina 1981) [=Triadotypus sodgianus, T. sodiques (l. c.)] NB01
    `--Discoidalia NB01
         |--Triadophlebiomorpha RP02
         `--Odonata [Stigmoptera] GE05
              |  i. s.: Epallagoidea PN04
              |           |--Epallage [Epallagidae] P74
              |           |--Litheuphaea carpenteri Fraser 1955 PN04
              |           |--Parazacallites aquisextana Nel 1988 PN04
              |           `--Zacallites balli Cockerell 1928 PN04
              |         Telebasis salva P74
              |--Panodonata [Anisozygoptera, Libellulina] RP02
              |    |--Euodonata RP02
              |    `--Tarsophlebioptera FB03
              |         |--Turanophlebia GE05
              |         `--Tarsophlebia FB03 [Tarsophlebiidae RP02]
              |              `--*T. eximia (Hagen 1862) FB03
              `--Protomyrmeleontoidea [Archizygoptera, Protozygoptera] RP02
                   |  i. s.: Lodevia longialata Nel et al. 1999 BN02
                   |--Kennedyidae RP02
                   |--Permolestidae RP02
                   |--Permepallage BN02 [Permepallagidae RP02]
                   |    `--P. augustissima Martynov 1938 BN02
                   |--Permagrionidae RP02
                   |--Protomyrmeleontidae RP02
                   `--Batkeniidae RP02

*Type species of generic name indicated

REFERENCES

[BN02] Béthoux, O., A. Nel, J. Lapeyrie, G. Gand & J. Galtier. 2002. Raphogla rubra gen. n., sp. n., the oldest representative of the clade of modern Ensifera (Orthoptera: Tettigoniidea, Gryllidea). European Journal of Entomology 99: 111-116.

[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.

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

[NB01] Nel, A., O. Bethoux, G. Bechly, X. Martínez-Delclòs & F. Papier. 2001. The Permo-Triassic Odonatoptera of the “protodonate” grade (Insecta: Odonatoptera). Annales de la Société Entomologique de France (N.S.) 37 (4): 501-525.

[P74] Paulson, D. R. 1974. Reproductive isolation in damselflies. Systematic Zoology 23 (1): 40-49.

[PN04] Petrulevičius, J. F. & A. Nel. 2004. A new damselfly family from the Upper Palaeocene of Argentina. Palaeontology 47: 109-116.

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

Stenophlebioptera

Specimen of Stenophlebia latreillei, copyright Günter Bechly.

Belongs within: Euepiproctophora.

The Stenophlebioptera are a group of fossil dragonflies known from the Lower Jurassic to the Lower Cretaceous (Fleck et al. 2003).

Synapomorphies (from Fleck et al. 2003): Nodal Cr very oblique, angle Cr-RA >140°; subnodus Sn very oblique, with or without cross-veins reaching it; CuA with broad area between two most distal posterior branches; straight supplementary longitudinal veins present in areas between IR2 and MP, near posterior wing margin.

<==Stenophlebioptera
    |--Liassostenophlebia Fleck, Bechly et al. 2003 [Liassostenophlebiidae] FB03
    |    `--*L. germanica Fleck, Bechly et al. 2003 FB03
    `--Stenophlebioidea FB03
         |--Prostenophlebia Nel & Martínez-Delclòs in Nel et al. 1993 [Prostenophlebiidae] FB03
         |    `--*P. jurassica Nel & Martínez-Delclòs in Nel et al. 1993 FB03
         `--Stenophlebiidae [Stenophlebiinae, Stenophlebinae] FB03
              |  i. s.: Sinostenophlebia Hong 1984 FB03
              |           `--S. zhangjiakowensis Hong 1984 FB03
              |--+--Cretastenophlebia Fleck, Bechly et al. 2003 FB03
              |  |    `--*C. mongolica Fleck, Bechly et al. 2003 FB03
              |  `--+--Hispanostenophlebia Fleck, Bechly et al. 2003 FB03
              |     |    `--*H. barremiana Fleck, Bechly et al. 2003 FB03
              |     `--Mesostenophlebia Fleck, Bechly et al. 2003 FB03
              |          `--*M. anglicana Fleck, Bechly et al. 2003 FB03
              `--Stenophlebia Hagen 1866 FB03
                   |  i. s.: S. casta (Hagen 1862) [=Heterophlebia casta] FB03
                   |         S. corami Nel & Jarzembowski 1996 FB03
                   |         S. lithographica (Giebel 1857) (see below for synonymy) FB03
                   |--*S. amphitrite (Hagen 1862) [=Heterophlebia amphitrite] FB03
                   |--S. eichstaettensis Nel et al. 1993 [=S. eichstattense] FB03
                   |--S. phryne (Hagen 1862) [=Heterophlebia phryne] FB03
                   `--+--S. karatavica Pritykina 1968 FB03
                      `--S. latreillei (Germar 1839) (see below for synonymy) FB03

Stenophlebia latreillei (Germar 1839) [=Agrion latreillei, Calopteryx latreillei, Heterophlebia latreillei; incl. H. aequalis Hagen 1862, Stenophlebia aequalis, Cordulegaster muensteri Hagen 1850, Diastatomma muensteri] FB03

Stenophlebia lithographica (Giebel 1857) [=Calopteryx lithographica, Heterophlebia lithographica] FB03

*Type species of generic name indicated

REFERENCES

[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.

Isophlebioptera

Bellabrunetia catherinae, copyright Ghedoghedo.

Belongs within: Euodonata.

The Isophlebioptera are a group of dragonflies known from the Triassic to Early Cretaceous of Eurasia. Most species are known only from wings, though the discovery of the more completely preserved Bellabrunetia catherinae from the Yixian Formation of China indicates that at least some exhibited a greater differentiation between fore and hind wings than modern dragonflies (Fleck & Nel 2002).

Synapomorphies (from Fleck & Nel 2002): Wings with vein RP3/4 slightly curved, more or less parallel to IR2, space between RP3/4 and MA distinctly expanded and transversed by several pectinated convex secondary branches of RP3/4; postdiscoidal space between MP and MA very narrow, primitively with only one row of cells between them; CuAa shortened, posterodistally indistinct (zig-zagged), distal space between MP and CuAa strongly expanded; all antenodal cross-veins between costal margin and ScP suppressed distad of Ax2 (reversed in some taxa). Hindwings with subdiscoidal space enlarged with bulged posterior margin, correlating with unique course of anal vein AA ('pseudo-anal loop').

<==Isophlebioptera
    |  i. s.: Italophlebia gervasuttii Whalley 1986 FN02
    |--Euthemis Pritykina 1968 [Euthemistidae] FN02
    `--+--Parazygoptera FN02
       |    |  i. s.: Cyclothemis [Cyclothemistidae] FN02
       |    |           `--C. sagulica Pritykina 1980 FN02
       |    |--Sphenophlebiidae FN02
       |    `--Euparazygoptera FN02
       |         |--Asiopteridae FN02
       |         `--Triassolestoidea FN02
       |              |--Triassolestes F71 [Progonophlebiidae, Triassolestidae FN02]
       |              |    `--T. epiphlebioides Tillyard 1918 F71
       |              `--Caraphlebia RP02 [Selenothemistidae FN02]
       |                   `--C. antarctica Carpenter 1969 RP02
       `--Isophlebiida FN02
            |  i. s.: Liadotypus Martynov 1937 [Liadotypidae] NB01
            |           `--*L. relictus Martynov 1937 NB01
            |--Archithemistidae FN02
            `--Isophlebioidea FN02
                 |--Isophlebiidae FN02
                 |    |--Anisophlebia Handlirsch 1908 FN02
                 |    |--Dinosamarura S02
                 |    `--Isophlebia Hagen 1866 FN02
                 |         `--I. aspasia Hagen 1866 FB03
                 `--Campterophlebiidae FN02
                      |--Bathmophlebia Pritykina 1970 FN02
                      |--Hypsomelana Pritykina 1968 FN02
                      |--Oreophlebia Pritykina 1970 FN02
                      |--Campterophlebia Bode 1905 FN02
                      |--Xantohypsa Pritykina 1970 FN02
                      |--Sogdophlebia Pritykina 1970 FN02
                      |--Hypsophlebia Pritykina 1970 FN02
                      |--Sagulia Pritykina 1970 FN02
                      |--Sarytashia Pritykina 1970 FN02
                      |--Gampsophlebia Pritykina 1980 FN02
                      |--Hypsothemis Pritykina 1968 FN02
                      |--Melanohypsa Pritykina 1968 FN02
                      |--Adelophlebia Pritykina 1980 FN02
                      |--Oshinia Pritykina 1985 FN02
                      |--Angaroneura Pritykina 1985 FN02
                      |--Dorsettia Whalley 1985 FN02
                      |--Karatawia Martynov 1925 [Karatawiidae] FN02
                      |--Pternopteron mirabile Pritykina 1970 FN02
                      |--Samarura gigantea RP02
                      |--Sibirioneura Pritykina 1985 FN02
                      |    `--S. amurica FN02
                      |--Bellabrunetia Fleck & Nel 2002 FN02
                      |    `--*B. catherinae Fleck & Nel 2002 FN02
                      `--Olonkia Pritykina 1985 FN02
                           `--O. elongata Pritykina 1985 FN02

*Type species of generic name indicated

REFERENCES

[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.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1-167.

[NB01] Nel, A., O. Bethoux, G. Bechly, X. Martínez-Delclòs & F. Papier. 2001. The Permo-Triassic Odonatoptera of the “protodonate” grade (Insecta: Odonatoptera). Annales de la Société Entomologique de France (N.S.) 37 (4): 501-525.

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

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

Euepiproctophora

Epiophlebia superstes, photographed by chochoensis.

Belongs within: Euodonata.
Contains: Stenophlebioptera, Anisoptera.

The odonate clade Euepiproctophora includes the living dragonflies and the Asian relict genus Epiophlebia. The name of the clade is derived from the development of the epiprocts at the end of the male's abdomen to form part of the claspers used in mating. Epiophlebia comprises two species of odonate found in Japan (E. superstes) and the Himalayas (E. laidlawi); though similar to a typical dragonfly in overall appearance, they have petiolate wings more similar to those of the damselflies in the Zygoptera (Grimaldi & Engel 2005).

See also: A halfway house, halfway down Honshu island.

Characters (from Fleck et al. 2003; Grimaldi & Engel 2005): Frons bulbous; thoracic interpleural suture dorsally present; epiprocts enlarged in male to form appendage of claspers; naiad with rectal chamber lined with respiratory pads; postdiscoidal area on wing broader than area between MA and RP3/4 along posterior wing margin.

<==Euepiproctophora
    |--Epiophlebia [Epiophlebiidae, Epiophlebioidea, Epiophlebioptera] FB03
    |    |--E. laidlawi GE05
    |    `--E. superstes (Selys 1889) FB03
    `--+--Erichschmidtia Pritykina 1968 [Erichschmidtiidae] FB03
       `--Anisopteromorpha [Heterophlebioidea] FB03
            |--Trigonoptera FB03
            |    |  i. s.: Gondvanogomphus Schlüter & Hartung 1982 [Gondvanogomphidae] FB03
            |    |--Stenophlebioptera FB03
            |    `--Anisoptera FB03
            `--Heterophlebioptera FB03
                 |--Myopophlebiidae FB03
                 |    |--Paraplagiophlebia loneuxi FB03
                 |    `--Paraheterophlebia marci Nel & Henrotay 1993 FB03
                 `--+--Liassophlebia [Liassophlebiidae] FB03
                    |    `--L. magnifica Tillyard 1925 FB03
                    `--+--Juraheterophlebia Fleck, Bechly et al. 2003 [Juraheterophlebiidae] FB03
                       |    `--*J. kazakhstanensis Fleck, Bechly et al. 2003 FB03
                       `--Heterophlebia [Heterophlebiidae] FB03
                            `--H. buckmani (Brodie 1849) FB03

*Type species of generic name indicated

REFERENCES

[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.

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

Triadophlebiomorpha

Diagram of preserved wing portion of Zygophlebiella curta, showing pattern of basal venation. From Nel et al. (2001).

Belongs within: Pandiscoidalia.

The Triadophlebiomorpha are a clade of Permian and Triassic insects of the odonate stem group, known from central Asia and Russia. Many triadophlebiomorphs have petiolate wings similar to those of modern damselflies; however, Xamenophlebia ornata is notable for its relatively broad wings with a strongly undulated hindwing.

Characters (from Nel et al. 2001): Wing with very long petiolus, with veins MP and Cu basally separated, distally fused together with AA, CuA + CuP + AA separate more distally from MP; MP distinctly curved distad of its origin at distal angle of discoidal cell; very long and oblique discoidal vein MAb; CuA without distal posterior branches.

<==Triadophlebiomorpha
    |  i. s.: Piroutetia Meunier 1907 [Piroutetiidae] NB01
    |           `--*P. liasina Meunier 1907 NB01
    |--Triadophlebiida NB01
    |    |--Mitophlebia [Mitophlebiidae] NB01
    |    `--Triadophlebioidea NB01
    |         |--Paurophlebia Pritykina 1981 [Paurophlebiidae] NB01
    |         |    `--P. lepida NB01
    |         `--Triadophlebiidae NB01
    |              |--Triadophlebia Pritykina 1981 NB01
    |              `--Cladophlebia Pritykina 1981 NB01
    |                   `--C. parvula Pritykina 1981 NB01
    `--Zygophlebiida NB01
         |--Xamenophlebioidea NB01
         |    |--Xamenophlebia Pritykina 1981 [Xamenophlebiidae] NB01
         |    |    `--*X. ornata Pritykina 1981 NB01
         |    `--Permophlebia Nel, Bethoux et al. 2001 [Permophlebiidae] NB01
         |         `--*P. uralica Nel, Bethoux et al. 2001 NB01
         `--Zygophlebiidae [Zygophlebioidea] NB01
              |--Zygophlebia Pritykina 1981 NB01
              |    `--*Z. ramosa Pritykina 1981 NB01
              |--Zygophlebiella Pritykina 1981 NB01
              |    `--*Z. curta Pritykina 1981 NB01
              |--Mixophlebia Pritykina 1981 NB01
              |    `--*M. mixta Pritykina 1981 NB01
              `--Cyrtophlebia Pritykina 1981 NB01
                   `--*C. sinuosa Pritykina 1981 NB01

*Type species of generic name indicated

==References==

[NB01] Nel, A., O. Bethoux, G. Bechly, X. Martínez-Delclòs & F. Papier. 2001. The Permo-Triassic Odonatoptera of the “protodonate” grade (Insecta: Odonatoptera). Annales de la Société Entomologique de France (N.S.) 37 (4): 501-525.