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).

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.: Palaeaeschna S02
    |         Juragomphus Nel, Bechly et al. 2001 [Juragomphidae] NB01
    |           `--*J. karatauensis Nel, Bechly et al. 2001 NB01
    |         Libellulium longialata GE05
    |--Petaluroidea [Petalurida] GE05
    |    |--Protolindeniidae RP02
    |    |--Aktassiinae [Aktassiidae] RP02
    |    |    |--Aktassia Pritykina 1968 NB01
    |    |    `--Aeschnogomphus Handlirsch 1906 NB01
    |    `--Petaluridae TH06
    |         |--Uropetala carovei (White 1846) HL02
    |         |--Phenes raptor Rambur 1842 FN02
    |         `--Petalura TH06
    |              |--P. gigantea Leach 1815 TH06
    |              |--P. hesperia Watson 1958 TH06
    |              |--P. ingentissima Tillyard 1908 TH06
    |              |--P. litorea Theischinger 1999 TH06
    |              `--P. pulcherrima Tillyard 1913 TH06
    `--Euanisoptera [Exophytica] RP02
         |--Gomphoidea GE05
         `--+--Cavilabiata GE05
            `--Aeshnoidea [Aeshnoptera] GE05
                 |  i. s.: Chrysogomphus Ren 1994 FB03
                 |           `--*C. beipiaoensis Ren 1994 FB03
                 |--Eumorbaeschnidae RP02
                 |--Telephlebiidae WO91
                 |--Mesuropetala RJ93 [Mesuropetalidae RP02]
                 |    `--M. koehleri RJ93
                 |--Cymatophlebia W13 [Cymatophlebiidae RP02]
                 |    `--C. longialata W13
                 |--Paramesuropetala [Liupanshaniidae] NB01
                 |    `--P. gigantea Bechly et al. 2001 NB01
                 `--Aeshnidae [Aeschnidae] TH06
                      |  i. s.: Gomphoaeschna SL89
                      |         *Tetracanthagyna plagiata (Waterhouse 1878) [=Gynacantha plagiata] SL89
                      |         Adversaeschna brevistyla (Rambur 1842) TH06
                      |         Boyeria A07
                      |         Basiaeshna A07
                      |         Oligoaeschna Selys Longchamps 1889 SL89
                      |           |--*O. modiglianii Selys Longchamps 1889 SL89
                      |           `--O. kunigamiensis (Ishida 1972) I92
                      |         Karschia Förster 1900 non Körb. 1865 (ICBN) T08
                      |           |--K. angulata [=Gynacantha angulata] T08
                      |           `--K. cornuta [=Gynacantha cornuta] T08
                      |         Acanthagyna dravida LM72
                      |--Anactinae WO91
                      |    |--Hemianax papuensis (Burmeister 1839) WO91, TH06 [=Anax papuensis HL02]
                      |    |--Anaciaeschna Selys 1878 WO91, SL89
                      |    |    `--A. jaspidea (Burmeister 1839) TH06 (see below for synonymy)
                      |    `--Anax TH06
                      |         |--A. amazili PP72
                      |         |--A. ephippiger [=Hemianax ephippiger] HL02
                      |         |--A. georgius Selys 1872 TH06
                      |         |--A. gibbosulus Rambur 1842 TH06
                      |         |--A. guttatus (Burmeister 1839) TH06
                      |         |--A. imperator RD77
                      |         `--A. junius RD77
                      |--Gynacanthaginae WO91
                      |    |--Agyrtacantha dirupta (Karsch 1889) WO91, TH06
                      |    |--Austrogynacantha Tillyard 1908 WO91, T08
                      |    |    `--*A. heterogena Tillyard 1908 [=Gynacantha heterogena de Selys ms] T08
                      |    `--Gynacantha WO91
                      |         |--G. dobsoni Fraser 1951 TH06
                      |         |--G. kirbyi Krüger 1898 TH06
                      |         |--G. mocsaryi Förster 1898 TH06
                      |         |--G. nourlangie Theischinger & Watson 1991 TH06
                      |         |--G. rosenbergi Kaup 1867 TH06
                      |         `--G. subinterrupta SL89
                      `--Aeshninae WO91
                           |--Caliaeschna T08
                           `--Aeschna GD00
                                |--A. brevistyla Rambur 1842 HL02
                                |--A. canadensis A07
                                |--A. crenata K01
                                |--A. cyanea GD00
                                |--A. forcipata L02
                                |--A. grandis K01
                                |--A. juncea K01
                                |--A. larvata S02
                                |--A. nigroflava I92
                                |--A. pryeri K00
                                |--A. sitchensis B-PE07
                                |--A. subarctica Walker 1906 I92
                                `--A. thomassoni Kirby 1900 K00

Anaciaeschna jaspidea (Burmeister 1839) TH06 [=Aeschna jaspidea SL89; incl. Anax taitensis SL89]

*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 10^th 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 2^a, 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 2^nd 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).

Last updated: 29 May 2022.

Zygoptera

Tropical rockmaster Diphlebia euphoeoides, copyright Wildiaries.

Belongs within: Euodonata.
Contains: Calopterygoidea, Lestoidea, Isostictidae, Coenagrionidae.

The Zygoptera, damselflies, are a group of generally more delicate odonates with similar, often petiolate fore- and hind wings. Damselflies also have broad heads with widely separated eyes, a strongly oblique thorax allowing the wings to be closed towards the rear of the body, and nymphs with three terminal gills (Grimaldi & Engel 2005). The basal division within the damselflies is between the clades Caloptera and Euzygoptera. The forewings possess a number of antenodal crossveins in the costal space in members of the Caloptera but the number of antenodal veins is reduced (usually to two) in Euzygoptera (Theischinger & Hawking 2006). The Caloptera are supported as a clade by basal recession of the midfork and an obsolete pterostigmal brace vein. The Eucaloptera, including most Caloptera except the Oligocene–Miocene European family Sieblosiidae, are also supported by their rectangular discoidal cell. Members of the clade Amphipterygida lack secondary antenodal crossveins between ScP and RA distal of the second primary antenodal Ax2 and also lack crossveins in at least the basal half of the antesubnodal space (Petrulevičius & Nel 2004).

Within the Euzygoptera, the Coenagrionoidea are characterised by an acute discoidal cell and short, braced pterostigma with wrinkled micro-sculpturing (Bechly 2012). Hemiphlebia mirabilis is a tiny, metallic green damselfly found in south-eastern Australia that differs from other damselflies in having an open discoidal cell (lacking an arculus) and a CuP vein that is almost straight in the first cell past the discoidal cell (Theischinger & Hawking 2006).

Characters (from Petrulevičius & Nel 2004): Wings with terminal kinks reduced in both CP and nodal furrow; tubular sclerotised canal of ScP along venter of postnodal costal margin obliterated; postnodal crossveins very numerous and dense; discoidal cell quadrangular, not divided into triangle and hypertriangle; vein RP2 distally displaced far from subnodus; CuA without secondary branching into CuAa and CuAb; oblique vein absent.

<==Zygoptera [Calopterygina]
    |  i. s.: Cretachistigma Jarzembowski et al. 1998 PN04
    |           |--C. essweini Bechly 1998 PN04
    |           `--C. greenwoodi Jarzembowski et al. 1998 PN04
    |         Paraphlebia Hagen 1861 PN04
    |         Philogenia Selys 1862 PN04
    |         Archilestes RD77
    |           |--A. californica D72
    |           `--A. grandis RD77
    |--Caloptera PN04
    |    |--Stenolestes RJ93 [Sieblosiidae PN04]
    |    |    |--S. fischeri Nel 1986 RJ93
    |    |    `--S. hispanicus Nel 1991 RJ93
    |    `--Eucaloptera PN04
    |         |--Calopterygoidea PN04
    |         `--Amphipterygida PN04
    |              |--Diphlebiidae PN04
    |              |    |--Philoganga [Philogangidae, Philoganginae] TH06
    |              |    `--Diphlebia [Diphlebiinae] TH06
    |              |         |--D. coerulescens Tillyard 1913 TH06 [=D. euphaeoides coerulescens C70]
    |              |         |--D. euphoeoides Tillyard 1907 TH06
    |              |         |--D. hybridoides Tillyard 1912 TH06
    |              |         |--D. lestoides (Selys 1853) TH06
    |              |         `--D. nymphoides Tillyard 1912 TH06
    |              `--Amphipterygoidea [Amphipterygida] PN04
    |                   |--Protamphipteryx Cockerell 1920 [Amphipterygidae] PN04
    |                   |--Latibasalia Petrulevičius & Nel 2004 [Latibasaliidae] PN04
    |                   |    |--*L. elongata Petrulevičius & Nel 2004 PN04
    |                   |    `--L. quispeae Petrulevičius & Nel 2004 PN04
    |                   `--Thaumatoneuridae [Thaumatoneuroidea] PN04
    |                        |--Thaumatoneurinae PN04
    |                        |    |--Thaumatoneura McLachlan 1897 PN04
    |                        |    `--Euarchistigma Carle & Wighton 1990 PN04
    |                        |         `--E. atrophium Carle & Wighton 1990 RJ93
    |                        `--Dysagrioninae PN04
    |                             |--Congqingia Zhang 1992 [Congqingiidae] PN04
    |                             |--Dysagrion Scudder 1878 PN04
    |                             |--Phenacolestes Cockerell 1908 PN04
    |                             `--Petrolestes Cockerell 1927 PN04
    |                                  `--P. hendersoni RJ93
    `--Euzygoptera GE05
         |--+--Lestoidea GE05
         |  `--Hemiphlebia [Hemiphlebiidae, Hemiphlebioidea] GE05
         |       `--H. mirabilis Selys 1869 TH06
         `--+--Lestoidea TH06 [Lestoideidae, Lestoideoidea GE05]
            |    |--L. barbarae Watson 1967 TH06
            |    |--L. brevicauda Theischinger 1996 TH06
            |    |--L. conjuncta Tillyard 1913 TH06
            |    `--L. lewisiana Theischinger 1996 TH06
            `--Coenagrionoidea [Coenagrionomorpha] GE05
                 |--Isostictidae WO91
                 |--Platystictidae WO91
                 |--Coenagrionidae TH06
                 |--Megaloprepus GE05 [Pseudostigmatidae WO91]
                 |    `--M. coerulatus RD77
                 |--Platycnemididae [Platycnemidae] FB03
                 |    |--Platycnemis E12
                 |    |    |--P. dealbata P74
                 |    |    |--P. pennipes P74
                 |    |    `--P. subdilatata de Selys 1847 E12
                 |    `--Coeliccia FB03
                 |         |--C. cyanomelas Ris 1912 FB03
                 |         |--C. flavicauda I92
                 |         |    |--C. f. flavicauda I92
                 |         |    `--C. f. masakii Asahina 1951 I92
                 |         `--C. ryukyuensis Asahina 1951 I92
                 |              |--C. r. ryukyuensis I92
                 |              `--C. r. amamii Asahina 1962 I92
                 `--Protoneuridae [Protoneurinae] RP02
                      |--Protonevra SL89
                      |--Notoneura O70
                      |--Eoprotoneura hyperstigma Carle & Wighton 1990 RJ93
                      `--Nososticta TH06
                           |--N. baroalba Watson & Theischinger 1984 TH06
                           |--N. coelestina (Tillyard 1906) TH06
                           |--N. fraterna (Lieftinck 1933) TH06
                           |--N. kalumburu Watson & Theischinger 1984 TH06
                           |--N. koolpinyah Watson & Theischinger 1984 TH06
                           |--N. koongarra Watson & Theischinger 1984 TH06
                           |--N. liveringa Watson & Theischinger 1984 TH06
                           |--N. mouldsi Theischinger 2000 TH06
                           |--N. pilbara Watson 1969 TH06
                           |--N. solida (Hagen 1860) TH06
                           |--N. solitaria (Tillyard 1906) TH06
                           `--N. taracumbi Watson & Theischinger 1984 TH06

*Type species of generic name indicated

REFERENCES

[C70] CSIRO. 1970. The Insects of Australia: A textbook for students and research workers. Melbourne University Press.

[D72] Davies, T. W. 1972. Oviposition of the California damsel fly, Archilestes californica MacLachlan. Pan-Pacific Entomologist 48 (1): 68–69.

[E12] Evenhuis, N. L. 2012. Publication and dating of the Exploration Scientifique de l'Algérie: Histoire Naturelle des Animaux Articulés (1846–1849) by Pierre Hippolyte Lucas. Zootaxa 3448: 1–61.

[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.

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

[O70] O’Farrell, A. F. 1970. Odonata (dragonflies and damselflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 241–261. Melbourne University Press.

[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: 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 10^th 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 2^a, 7: 444–484.

[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: 30 May 2022.

Libellulidae

Male Rhyothemis plutonia, from here.

Belongs within: Paneurypalpidomorpha.
Contains: Brachydiplax, Orthetrum, Libellula, Agrionoptera.

The Libellulidae, skimmers and perchers, are a diverse group of dragonflies with antenodal costal and subcostal veins in strict alignment, and a rounded base to the hindwing in both sexes (Theischinger & Hawking 2006). Members of the group vary widely in appearance; some have strongly patterned wings whereas others have entirely transparent wing membranes. The family is divided between several subfamilies, primarily on the basis of wing venation characters. Members of the largest subfamily, the Sympetrinae, have fore wings with the last antenodal crossvein not extending beyond the subcosta, the arculus between the first and second antenodal crossveins, and distinct radial and medial planates, and hind wings with a broad base and a highly visible anal loop. The Libellulinae usually have the arculus at or distal to the second antenodal crossvein in both pairs of wings; it is always so at least in the hind wing. The Pantalinae are a subfamily of broad-winged, strong fliers (Watson & O'Farrell 1991); the pantropical Pantala flavescens may be the single most widespread odonatan species.

See also: Libellulidae: on the wing.

Characters (from Theischinger & Hawking 2006): Wings with antenodal costal and subcostal veins in strict alignment, primary antenodals not discernible; sectors of arculus generally stalked at base; anal loop generally well developed; hindwing base rounded in both sexes. Lavae with labium ladle-shaped, bearing premental and palpal setae; labial palps smooth to strongly toothed; abdomen generally laterally and (more rarely) dorsally armed.

<==Libellulidae
    |--Trithemistinae WO91
    |    |--Austrothemis nigrescens (Martin 1901) WO91, TH06
    |    `--Trithemis SL89
    |         |--T. adelpha Selys 1878 SL89
    |         |--T. festiva [=Libellula festiva; incl. Dythemis infernalis] SL89
    |         |--T. fraterna SL89
    |         `--T. tirbyi Im92
    |--Zyxommatinae WO91
    |    |--Tholymis tillarga (Fabricius 1798) WO91, TH06
    |    `--Zyxomma TH06
    |         |--*Z. petiolatum Rambur 1842 SL89, TH06
    |         |--Z. atlanticum SL89
    |         |--Z. elgneri Ris 1913 TH06
    |         |--Z. multinervorum Carpenter 1897 TH06
    |         `--Z. obtusum SL89
    |--Brachydiplactinae WO91
    |    |--Brachydiplax TH06
    |    |--Raphismia bispina (Hagen 1867) WO91, TH06
    |    `--Nannophya Rambur 1842 WO91, T08a
    |         |--N. australis Brauer 1865 TH06
    |         |--N. dalei (Tillyard 1908) TH06
    |         |--N. occidentalis (Tillyard 1908) TH06 [=N. dalei occidentalis C70]
    |         |--N. paulsoni Theischinger 2003 TH06
    |         `--N. pygmaea [incl. N. exigua] SL89
    |--Tetrathemistinae WO91
    |    |--Tetrathemis TH06
    |    |    |--T. flavescens T08c
    |    |    |--T. irregularis TH06
    |    |    |    |--T. i. irregularis TH06
    |    |    |    `--T. i. cladophila Tillyard 1908 TH06
    |    |    `--T. platyptera TH06
    |    `--Nannophlebia WO91
    |         |--N. eludens Tillyard 1908 TH06
    |         |--N. imitans T08c
    |         |--N. injibandi Watson 1969 TH06
    |         |--N. lorquini T08c
    |         |--N. mudginberri Watson & Theischinger 1991 TH06
    |         `--N. risi Tillyard 1913 TH06
    |--Pantalinae WO91
    |    |--Pantala flavescens (Fabricius 1798) TH06 [incl. Libellula terminalis SL89, L. viridula SL89]
    |    |--Hydrobasileus brevistylus (Brauer 1865) WO91, TH06
    |    |--Trapezostigma WO91
    |    |    |--T. loewi Brauer 1866 T13
    |    |    `--T. stenoloba PH10
    |    |--Camacinia WO91
    |    |    |--C. gigantea TH06
    |    |    `--C. othello Tillyard 1908 TH06
    |    `--Tramea WO91
    |         |--T. euryale SL89
    |         |--T. eurybia Selys 1878 TH06
    |         |--T. loewii Kaup 1866 TH06
    |         |--T. propinqua Lieftinck 1942 TH06
    |         `--T. stenoloba (Watson 1962) TH06
    |--Rhyothemis TH06 [Rhyothemistinae WO91]
    |    |--R. alcestis T08b
    |    |--R. braganza Karsch 1890 TH06
    |    |--R. chloe T08b
    |    |--R. curiosa Selys Longchamps 1889 (see below for synonymy) SL89
    |    |--R. fulgens Selys Longchamps 1889 SL89
    |    |--R. fuliginosa SL89
    |    |--R. graphiptera (Rambur 1842) TH06
    |    |--R. phyllis (Sulzer 1776) TH06
    |    |    |--R. p. phyllis TH06
    |    |    |--R. p. beatricis TH06
    |    |    `--R. p. chloe TH06
    |    |--R. plutonia SL89
    |    |--R. princeps Kirby 1894 TH06
    |    |--R. resplendens Selys 1878 TH06
    |    `--R. vidua SL89
    |--Libellulinae WO91
    |    |--Orthetrum WO91
    |    |--Notolibellula bicolor Theischinger & Watson 1977 WO91, TH06
    |    |--Libellula RS10
    |    |--Agrionoptera WO91
    |    |--Potamarcha congener (Rambur 1842) O70, TH06
    |    |--Nannodythemis Brauer 1868 T08a
    |    |    |--*N. australis Brauer 1868 T08a
    |    |    |--N. dalei Tillyard 1908 T08a
    |    |    `--N. occidentalis Tillyard 1908 T08a
    |    |--Lathrecista WO91
    |    |    |--L. asiatica TH06
    |    |    |    |--L. a. asiatica TH06
    |    |    |    `--L. a. festa (Selys 1879) TH06 [=Agrionoptera festa SL89, Orthemis festa SL89]
    |    |    |--L. difficilis [=Agrionoptera difficilis, Orthemis difficilis] SL89
    |    |    |--L. pectoralis [=Agrionoptera pectoralis, Orthemis pectoralis] SL89
    |    |    `--L. simulans [=Agrionoptera simulans, Orthemis simulans] SL89
    |    |--Zygonidia Kirby 1900 K00a
    |    |    `--*Z. insignis Kirby 1900 K00a
    |    `--Acisoma K00a
    |         |--A. ascalaphoides SL89
    |         `--A. panorpoides (Ramb. 1842) [=Libellula panorpoides] K00a
    `--Sympetrinae WO91
         |--Nannodiplax rubra Brauer 1868 WO91, TH06
         |--Rhodothemis lieftincki Fraser 1954 WO91, TH06
         |--Crocothemis WO91
         |    |--C. nigrifrons (Kirby 1894) TH06
         |    |--C. servilia (Drury 1773) [=Libellula servilia] K00a
         |    `--C. soror [=Trithemis soror] SL89
         |--Sympetrum K01
         |    |--S. corruptum [=Tarnetrum corruptum] A72
         |    |--S. flaveolum K01
         |    |--S. internum A07
         |    |--S. obtrusum A07
         |    |--S. pedemontanum K01
         |    |--S. rubicundulum A07
         |    |--S. sanguineum K01
         |    |--S. scoticum K01
         |    `--S. striolatum RD77
         |--Diplacodes Kirby 1889 WO91, FB03
         |    |--*D. tetra K00a [=Libellula tetra SL89, Diplacina tetra SL89]
         |    |--D. bipunctata (Brauer 1865) TH06
         |    |--D. concinna [=Libellula concinna, Diplacina concinna] SL89
         |    |--D. flavistyla [=Libellula flavistyla, Diplacina flavistyla] SL89
         |    |--D. haematodes (Burmeister 1839) TH06
         |    |--D. melanopsis (Martin 1901) TH06
         |    |--D. nebulosa (Fabricius 1793) TH06 [=Libellula nebulosa SL89, Diplacina nebulosa SL89]
         |    `--D. trivialis (Rambur 1842) TH06 [=Libellula trivialis K00a, Trithemis trivialis K00a]
         `--Neurothemis WO91 [incl. Polynevra SL89]
              |--N. basalis SL89
              |--N. degener Selys Longchamps 1879 [incl. Libellula intermedia] SL89
              |--N. feralis [incl. Libellula equestris] SL89
              |--N. fluctuans Selys 1879 SL89
              |    |--N. f. fluctuans SL89
              |    `--N. f. disparilis Kirby 1889 SL89
              |--N. oligoneura Brauer 1867 TH06
              |--N. stigmatizans (Fabricius 1775) TH06
              `--N. tullia (Drury 1773) [=Libellula tullia] K00a

Libellulidae incertae sedis:
  Diastatops NB01
  Nannothemis Brauer 1868 FB03
  Palaeothemis tillyardi Fraser 1923 FB03
  Neodythemis Karsch 1889 FB03
  Diplax trivialis SL89
  Tyriobapta torrida SL89
  Lepthemis vesiculosa SL89
  Huonia melvillensis Brown & Theischinger 1998 TH06
  Celithemis eponina A07
  Ladona P74
  Erythemis rufa A07, T08c
  Pachydiplex A07
  Perithemis A07
  Orchithemis [incl. Calothemis] SL89
    |--O. pruinans (Selys 1878) [=Calothemis pruinans] SL89
    `--O. pulcherrima Brauer 1878 [incl. Calothemis exsudans Selys 1878] SL89
  Nesciothemis Longfield 1955 L55
    |--*N. farinosum (Förster 1898) (see below for synonymy) L55
    `--N. fitzgeraldi (Pinhey 1955) [=Orthetrum fitzgeraldi] L55
  Orthemis SL89
    |--O. congener SL89
    `--O. lineata SL89
  Libella SL89
    |--L. clelia SL89
    |--L. glauca Brauer 1865 SL89
    |--L. luzonica Brauer 1868 SL89
    `--L. pruinosa [=Libellula pruinosa] SL89
  Diplacina SL89
    |--*D. nana SL89
    |--D. braueri SL89
    |--D. militaris SL89
    |--D. nigrilabris SL89
    |--D. obscura SL89
    `--D. smaragdina SL89
  Leucorrhinia K01
    |--L. frigida MG06
    |--L. hudsonica B-PE07
    |--L. intacta MG06
    |--L. intermedia Iw92
    |    |--L. i. intermedia Iw92
    |    `--L. i. ijimai Asahina 1961 Iw92
    |--L. pectoralis K01
    |--L. proxima MG06
    `--L. rubicunda K01
  Condalia woottoni Whalley & Jarzembowski 1985 RJ93
  Zygonyx Hagen 1867 K00a, K00b
    `--*Z. iris K00b
  Neurocena Kirby 1900 K00b
    `--*N. ida [=Zygonyx ida; incl. Pseudomacromia luxuriosa Karsch 1893] K00b
  Schizonyx Karsch 1890 K00b
    `--S. luctifera (De Selys 1869) [=Zygonyx luctifera] K00b
  Cratilla Kirby 1900 K00b
    `--*C. metallica (Brauer 1878) [=Orthemis metallica, Protorthemis metallica] K00b
  Nesoxenia K00b

*Nesciothemis farinosum (Förster 1898) [=Orthetrum farinosum; incl. Hadrothemis hyalina Fraser 1927 (n. n.), O. pollinosum Karsch 1899] L55

Rhyothemis curiosa Selys Longchamps 1889 [incl. R. curiosa var. apicalis Selys Longchamps 1889, R. curiosa var. transversa Selys Longchamps 1889] SL89

*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.

[A72] Arnaud, P. H., Jr. 1972. Mass movement of Sympetrum corruotum (Hagen) (Odonata: Libellulidae) in central California. Pan-Pacific Entomologist 48 (1): 75–76.

[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.

[C70] CSIRO. 1970. The Insects of Australia: A textbook for students and research workers. Melbourne University Press.

[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.

[Im92] Imes, R. 1992. The Practical Entomologist. Aurum Press: London.

[Iw92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[K00a] Kirby, W. F. 1900a. 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.

[K00b] Kirby, W. F. 1900b. On the species which have been included in Zygonyx, Hagen and De Selys. Annals and Magazine of Natural History, series 7, 5: 539–542.

[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.

[L55] Longfield, C. 1955. The Odonata of N. Angola. Part I. Diamang Publicações Culturais 27: 11–63.

[MG06] Mallatt, J., & G. Giribet. 2006. Further use of nearly complete 28S and 18S rRNA genes to classify Ecdysozoa: 37 more arthropods and a kinorhynch. Molecular Phylogenetics and Evolution 40: 772–794.

[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.

[O70] O’Farrell, A. F. 1970. Odonata (dragonflies and damselflies). In: CSIRO. The Insects of Australia: A textbook for students and research workers pp. 241–261. Melbourne University Press.

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

[PH10] Pinder, A. M., S. A. Halse, R. J. Shiel & J. M. McRae. 2010. An arid zone awash with diversity: patterns in the distribution of aquatic invertebrates in the Pilbara region of Western Australia. Records of the Western Australian Museum Supplement 78 (1): 205–246.

[RS10] Regier, J. C., J. W. Shultz, A. Zwick, A. Hussey, B. Ball, R. Wetzer, J. W. Martin & C. W. Cunningham. 2010. Arthropod relationships revealed by phylogenomic analysis of nuclear protein-coding sequences. Nature 463: 1079–1083.

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th 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 2^a, 7: 444–484.

[T13] Taylor, C. K. 2013. Annotated bibliography for Barrow Island terrestrial invertebrates. Records of the Western Australian Museum Supplement 83: 135–144.

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

[T08a] Tillyard, R. J. 1908a. On the genus Nannodythemis, with descriptions of new species. [Neuroptera: Odonata]. Proceedings of the Linnean Society of New South Wales 33: 444–455, pl. 6.

[T08b] Tillyard, R. J. 1908b. Notes and exhibits. Proceedings of the Linnean Society of New South Wales 33: 629.

[T08c] Tillyard, R. J. 1908c. On some remarkable Australian Libellulinae. Part II. Descriptions of new species. Proceedings of the Linnean Society of New South Wales 33: 637–649, pl. 14.

[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: 28 May 2022.

Calopterygoidea

Male yellow bush dart Psilocnemis marginipes, copyright Jee & Rani Nature Photography.

Belongs within: Zygoptera.
Contains: Rhinocypha.

The Calopterygoidea are a group of damselflies characterised by broad, often colourful wings. In members of the family Calopterygidae, the wings are densely reticulate with numerous cross- and supplementary longitudinal veins, and have non-petiolate bases. Historically, this family has been recognised as the Agrionidae or Agriidae, a name that has also been used to encompass many, or all, of the damselflies. Modern classifications do not include this family with the type genus Agrion no longer being treated as a valid name. Other families include the Chlorocyphidae, an Old World tropical group with a narrow head, close eyes, and a produced, snout-like clypeus (Richards & Davies 1977).

<==Calopterygoidea [Agrioidea, Calopterygomorpha]
    |--Eosagrionidae FB03
    |--Dicteriastidae WO91
    |--Pseudolestes Kirby 1900 K00 [Pseudolestidae RD77]
    |    `--*P. mirabilis Kirby 1900 K00
    |--Heliocharitidae RD77
    |--Polythoridae WO91
    |    |--Euthore fasciata (Hagen 1853) FB03
    |    `--Cora ZP01
    |         |--C. chiribiquete Zloty & Pritchard 2001 ZP01
    |         |--C. munda ZP01
    |         `--C. terminalis ZP01
    |--Euphaeidae WO91
    |    |--Epallagites avus RJ93
    |    `--Euphaea SL89
    |         |--E. aspasia SL89
    |         |--E. bocki MacLachl. 1880 SL89
    |         |--E. variegata SL89
    |         `--E. yayeyamensis I92
    |--Chlorocyphidae [Libellaginidae] FB03
    |    |--Libellago Selys 1840 FB03
    |    |--Heliocypha fenestrata TH06
    |    `--Rhinocypha TH06
    |--Calopterygidae [Calopteryginae] W75
    |    |--Psolodesmus mandarinus McLachlan 1870 FB03
    |    |--Neurobasis TH06
    |    |    |--N. australis Selys 1897 TH06
    |    |    `--N. chinensis TH06
    |    |--Mnais FB03
    |    |    |--M. andersoni McLachlan 1873 FB03
    |    |    |--M. nawai TMS91
    |    |    `--M. pruinosa TMS91
    |    |--Micromerus Rambur 1842 SL89, BP02
    |    |    |--M. aurantiacus SL89
    |    |    |--M. snellemani SL89
    |    |    `--M. sumatranus SL89
    |    |--Calopteryx W75
    |    |    |--C. aequabilis W75
    |    |    |--C. maculata W75
    |    |    |--C. splendens W75
    |    |    `--C. virgo W75
    |    |--Vestalis SL89
    |    |    |--V. amaena SL89
    |    |    |--V. luctuosa SL89
    |    |    |--V. lugens [incl. V. coracina] SL89
    |    |    `--V. melania SL89
    |    `--Dysphaea Selys 1853 SL89, PN04
    |         |--D. dimidiata SL89
    |         |--D. limbata SL89
    |         |--D. lugens SL89
    |         `--D. sublimbata SL89
    `--Agriidae [Agrionidae, Agrioninae] RD77
         |--Onychargia atrocyana SL89
         |--Stenobasis oscillans SL89
         |--Allonevra insignis SL89
         |--Podolestes SL89
         |    |--P. chrysopus Selys Longchamps 1889 SL89
         |    `--P. orientalis SL89
         |--Trichocnemis membranipes [=Platycnemis membranipes] SL89
         |    |--T. m. membranipes SL89
         |    `--T. m. silenta SL89
         |--Disparonevra SL89
         |    |--D. notostigma SL89
         |    `--D. verticalis SL89
         |--Sympycna K01
         |    |--S. fusca K01
         |    `--S. paedisca K01
         |--Erythromma najas K01
         |    |--E. n. najas I92
         |    `--E. n. baicalensis Belyshev 1964 I92
         |--Agrion K01
         |    |--A. hastulatum K01
         |    |--A. hylas (Trybom 1889) I92
         |    |--A. pulchellum K01
         |    |--A. splendens RD77
         |    |--A. telluris S02
         |    `--A. virgo L02
         |--Psilocnemis SL89
         |    |--P. annulata SL89
         |    |    |--P. a. annulata SL89
         |    |    |--P. a. ciliata SL89
         |    |    |--P. a. serapica SL89
         |    |    `--P. a. subannulata SL89
         |    |--P. imbricata SL89
         |    `--P. marginipes [=Platycnemis marginipes, incl. Pl. lacteola, Psilocnemis striatipes] SL89
         |--Matrona basilaris Selys 1853 K00
         |    |--M. b. basilaris I92
         |    |--M. b. japonica Foerster 1897 I92
         |    `--M. b. nigripectus I92
         |--Bayadera brevicauda K00, I92
         |    |--B. b. brevicauda I92
         |    `--B. b. ishigakiana Asahina 1964 I92
         `--Pseudophaea decorata (de Selys 1853) [=Euphaea decorata] K00

*Type species of generic name indicated

REFERENCES

[BP02] Bologna, M. A., & J. D. Pinto. 2002. The Old World genera of Meloidae (Coleoptera): a key and synopsis. Journal of Natural History 36 (17): 2013–2102.

[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.

[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.

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

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

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms' General Textbook of Entomology 10^th 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 2^a, 7: 444–484.

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

[TMS91] Tsurusaki, N., M. Murakami & K. Shimokawa. 1991. Geographic variation of chromosomes in the Japanese harvestman, Gagrellopsis nodulifera, with special reference to a hybrid zone in western Honshu. Zoological Science 8: 265–275.

[W75] Waage, J. K. 1975. Reproductive isolation and the potential for character displacement in the damselflies Calopteryx maculata and C. aequabilis (Odonata: Calopterygidae). Systematic Zoology 24 (1): 24–36.

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

[ZP01] Zloty, J., & G. Pritchard. 2001. Cora chiribiquete spec. nov., a new damselfly species from Colombia (Zygoptera: Polythoridae). Odonatologica 30 (2): 227–232.

Last updated: 30 May 2022.

Diaphanopterodea

Reconstruction and photograph of hind wing of Diaphanoptera munieri, from Bethoux & Nel (2003).

Belongs within: Palaeoptera.

The Diaphanopterodea are a group of plant-sucking insects known from the Middle Carboniferous to the Early Permian, noteworthy for having evolved the ability to fold the wings back along the body apparently independently of the Neoptera. The earliest Prochoropteridae are known from the Westphalian of North America (Sinitshenkova 2002).

Characters (from Sinitshenkova 2002): Robust or slender insects of medium to large size (20-80 mm wingspread). Head small with convex lateral eyes, and short to medium length sucking beak directed below. Prothoracic paranota lateral, small. Wings similar, usually almost identical in form and venation of both pairs, lacking archedictyon, moderately or well costalised (with C, SC, and R thick and running relatively close to each other, though costal space sometimes rather wide), with alternating convex (C, R, MA, CuA, A) and concave (RS, MP, CuP), with costal margin straight to moderately convex, MA simple, tightened sub-basally by cross-vein to or fused for a distance with RS, M₅ usually retained, cross-vein-like or forming false CuA base. Legs with tarsi three-segmented, otherwise not evidently specialised. Abdomen segments lacking paranota. Ovipositor cutting, with sheath shorter than stylets, bearing long stylus. Cerci very long, very rarely found in males.

<==Diaphanopterodea [Diaphanopterida]
    |  i. s.: Philiasptilon BN03
    |           |--*P. maculosum Zalessky 1931 BN03
    |           `--P. hueneckeni Pinto & Pinto de Ornellas 1978 BN03
    |         Diaphterum Pinto 1998 BN03
    |--Biarmohymenidae S02
    |--Elmoidae BN03
    |--Martynoviidae S02
    |--Prochoroptera Handlirsch 1911 BN03 [Prochoropteridae S02]
    |    `--P. calopteryx RJ93
    |--Asthenohymen [Asthenohymenidae] S02
    |    `--A. uralicum S02
    |--Paruralia [Paruraliidae] S02
    |    `--P. rohdendorfi S02
    |--Permohymen [Permohymenidae] RJ93
    |    `--P. schucherti K-P91
    |--‘Uralia’ Kukalova-Peck & Sinichenkova 1992 S02, EH19 (see below for synonymy)
    |    |--U. maculata Kukalova-Peck & Sinitshenkova 1992 K03 [=Permuralia maculata S02]
    |    `--U. rodendorfi K-P91
    `--Diaphanoptera Brongniart 1893 (see below for synonymy) BN03
         |--*D. munieri Brongniart 1893 [incl. Diaphanoptera superba Meunier 1908, Diaphanopterites superbus] BN03
         |--D. scudderi Brongniart 1893 (n.d.) BN03
         `--D. vetusta Brongniart 1893 BN03

‘Uralia’ Kukalova-Peck & Sinichenkova 1992 nec Mulsant & Verreaux 1866 nec Likharev 1925 nec Martynova 1976 S02, EH19 [Parelmoidae BN03]

Diaphanoptera Brongniart 1893 [incl. Diaphanopterites Handlirsch 1919, Pseudanthracothremma Handlirsh 1906; Diaphanopteridae, Diaphanopteritidae] BN03

*Type species of generic name indicated

REFERENCES

[BN03] Bethoux, O., & A. Nel. 2003. Revision of Diaphanoptera species and new diagnosis of Diaphanopteridae (Palaeoptera: Diaphanopteridae). Journal of Paleontology 77 (5): 1016–1020.

[EH19] Énay, R., & M. K. Howarth. 2019. Part L, revised, volume 3B, chapter 7: Systematic descriptions of the Perisphinctoidea. Treatise Online 120: 1–184.

[K03] Kluge, N. J. 2003. About evolution and homology of genital appendages of insects. Trudy Russkogo Entomologicheskogo Obshestva [Proceedings of the Russian Entomological Society] 74: 3–16.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[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. Superorder Dictyoneuridea Handlirsch, 1906 (=Palaeodictyopteroidea). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 115–124. Kluwer Academic Publishers: Dordrecht.

Last updated: 26 May 2022.

Polyneoptera

Diagram of holotype of the Late Carboniferous Parahomalophlebia courtini, from Rasnitsyn (2002).

Belongs within: Pterygota.
Contains: Orthopteroidea, Perlidea.

The Polyneoptera is a major subdivision of the insects, including grasshoppers, cockroaches and related taxa. Though members of this group are often broadly similar in habitus, its monophyly remains uncertain.

Characters (from Rasnitsyn 2002, as Gryllones): Large to medium size, rarely small. Mouthparts chewing. Unless lost, wings resting flat over body (neither roof-like nor spread), with contralateral wings overlapping each other in part or completely, with hind wing anal area usually enlarged and, when folded, bent down along line running before 2A (occasionally before 1A), with common base of anal veins reverted upside down. Thoracic venter usually not cryptosternous. Third valve (gonocoxite 9 with or without its style) forming main intromittent part of ovipositor. Male gonostylus free or, usually, lost. Paracercus lost. Ovaries usually panoistic. Development embryonicised, eclosing nymph with compound eyes, ocelli and usually with full number of tarsomeres, and usually without adult moulting.

<==Polyneoptera (see below for synonymy)
    |  i. s.: Chresmodidae [Chresmododea] R02
    |           |--Gryllidium oweni R02
    |           |--Saurophthirodes R02
    |           `--Chresmoda Germar 1839 [incl. Propygolampis Weyenbergh 1874] R02
    |                |--C. aquatica R02
    |                |--C. obscura [incl. Propygolampis giganteus] R02
    |                `--C. orientalis R02
    |         Polyernus complanatus R02
    |--Thoronysis [Thoronysidae] R02
    |--Prototettix [Prototettigidae] R02
    |--Eucaenus [Eucaenidae] R02
    |    `--E. ovalis Scudder 1885 BN02
    |--Spanioderidae R02
    |    |--Propteticus R02
    |    |--Dieconeura R02
    |    `--Axiologus Handlirsch 1906 R02, BN02
    |         `--A. thoracicus Handlirsch 1906 BN02
    `--+--Parahomalophlebia [Homalophlebiidae] R02
       |    `--P. courtini R02
       `--+--Orthopteroidea TW05
          `--Perlidea TW05

Polyneoptera [Anartioptera, Cacurgida, Eoblattida, Gryllones, Orthopterodida, Paurometabola, Phasmatomorpha, Plecopterida, Plecopterodea, Polyorthoptera, Protoblattodea]

*Type species of generic name indicated

REFERENCES

[BN02] Béthoux, O., & A. Nel. 2002. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1-88.

[R02] Rasnitsyn, A. P. 2002. Infraclass Gryllones Laicharting, 1781. The grylloneans (=Polyneoptera Martynov, 1938). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 254-262. Kluwer Academic Publishers: Dordrecht.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240-257.

Megasecoptera

Reconstruction of Sylvohymen sibiricus by A. G. Ponomarenko, from Sinitshenkova (2002).

Belongs within: Pterygota.

The Megasecoptera is a group of plant-sucking insects with elongate, petiolate wings known from the Middle Carboniferous to Late Permian (Sinitshenkova 2002).

Characters (from Sinitshenkova 2002): Size medium to large (wing length 10-100 mm). Body slender (more robust in Eubleptoptera). Head small, prognathous, with large convex eyes and long sucking beak. Antenna about as long as body. Thoracic segments of similar size, pronotal paranota large in Eubleptoptera, reduced to several thorn-like projections or lost in others. Wings spread permanently, fore and hind wings alike, long, often petiolate, with fore margin straight (relatively short and wide in small, dipterous Permothemistidae). Cross-veins not numerous, often forming regular rows, RS and MA bases connected by stabilised cross-vein and usually angular there (unless cross-vein replaced by temporal fusion of both RS and MA for a distance, or cross-vein slanted and thus forming false base of either). Fore leg short, middle and hind legs poorly known. Thorax and abdomen sometimes with cuticular projections, long or short, simple or branched, and arranged in regular rows. Male genitalia with 2-3-segmented claspers, male cercus (where known) sinuate basally. Ovipositor (where known) short, wide. Nymph (where known) similar to adult but with beak short, wing pads bent backward when large, pterothoracic segments with lateral spines near wing pad base, abdominal terga with hind margin denticulate. Imaginal moults not proven.

<==Megasecoptera [Mischopterida]
    |  i. s.: Pseudohymen GE05
    |--Eubleptoptera [Eubleptidodea, Eubleptina, Eubleptoidea] GE05
    |    |--Anchineuridae GE05
    |    |--Engisopteridae GE05
    |    |--Sphecorydaloididae GE05
    |    |--Eubleptus S02 [Eubleptidae GE05]
    |    |    `--E. danielsi S02
    |    `--Namurodiapha S02 [Namurodiaphidae GE05]
    |         `--N. sippelorum S02
    `--+--Dicliptera [Archodonata, Permothemistida, Permothemistina] GE05
       |    |  i. s.: Permothemidia caudata D70
       |    |--Ogassidae S02
       |    |--Diathema S02 [Diathemidae GE05, Diathemoptera]
       |    |    `--D. tenerum S02
       |    |--Doter D70 [Doteridae S02]
       |    |    `--D. minor D70
       |    |--Rectineura D70 [Rectineuridae S02]
       |    |    `--R. lineata D70
       |    |--Kansasia S02 [Kansasiidae GE05]
       |    |    `--K. pulchra S02
       |    `--Permothemistidae S02
       |         |--Permothemis S02
       |         `--Ideliella S02
       |--Protohymenoptera [Aspidothoracina] GE05
       |    |--Aspidothoracidae GE05
       |    |--Aspidohymenidae GE05
       |    |--Brodiidae GE05
       |    |--Brodiopteridae GE05
       |    |--Protohymenidae GE05
       |    |--Caulopteridae GE05
       |    |--Karoohymen S02 [Scytohymenidae GE05]
       |    |    `--K. delicatulus S02
       |    |--Ancoptera S02 [Ancopteridae GE05]
       |    |    `--A. permiana S02
       |    |--Hana S02 [Hanidae GE05]
       |    |    `--H. filia S02
       |    `--Sylvohymen S02 [Bardohymenidae GE05]
       |         |--S. peckae S02
       |         |--S. robustus S02
       |         `--S. sibiricus S02
       `--Eumegasecoptera [Mischopterina] GE05
            |--Allectoneuridae S02
            |--Moravohymenidae GE05
            |--Carbonopteridae GE05
            |--Corydaloididae GE05
            |--Foririidae S02
            |--Sphecopteridae GE05
            |--Ichnoptilidae S02
            |--Raphidiopseidae S02
            |--Parabrodiidae S02
            |--Aykhal [Aykhalidae] S02
            |    `--A. helenae S02
            |--Vorkutia S02 [Vorkutiidae GE05]
            |    `--V. dimina S02
            `--Mischoptera [Mischopteridae] GE05
                 |--M. douglassi S02
                 `--M. nigra GE05

*Type species of generic name indicated

REFERENCES

[D70] Demoulin, G. 1970. Remarques critiques sur des larves «ephemeromorphes» du Permien. Bulletin de l’Institut Royal des Sciences Naturelles de Belgique 46 (3): 1-10.

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

[S02] Sinitshenkova, N. D. 2002. Superorder Dictyoneuridea Handlirsch, 1906 (=Palaeodictyopteroidea). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 115-124. Kluwer Academic Publishers: Dordrecht.