Isoperla grammatica, photographed by N. Sloth.

Belongs within: Perlidea.
Contains: Grypopterygomorpha.

The Plecoptera, stoneflies, are a group of mostly detritivorous or omnivorous insects with nymphs that are mostly aquatic or, if terrestrial, found in damp habitats.

Characters (from Sinitshenkova 2002a): Body size moderately small to large. Head prognathous, with mouthparts chewing (mandibles sometimes much reduced). Thorax structure little modified, prothorax large, lacking wide paranota, meso- and metathoraces of similar structure. Legs cursory (with swimming hairs in many nymphs), with tarsus 3-segmented. Wings membranous, almost completely overlapping each other over abdomen at rest, with pre-anal venation similar in both pairs, excluding RS which lost its base and originated from M in hind wing (nymphal wing pad with RS base preserved), and CuA which is usually (primarily) simple in hind wing; hind wing anal area usually large. Wing venation not especially rich, with main veins branching at or beyond wing midlength, more often forked, RS and CuA up to 4 or 5 branches (in ground plan CuA is similar to M in having 2 branches only). Cross veins not numerous besides 2 characteristic rows aligned CuA, with most constant r-rs and r-m near RS and M forkings (roughly at level of SC apex). RS typically and probably plesiomorphically originating near wing base and running parallel to R. Flight functionally four-winged, in-phase. mainly anteromotoric in spite hind wing widened basally. Abdomen with full number of segments, lacking paranota. Primary ovipositor lost. Cerci always present, usually long, multisegmented, rarely one-segmented in adult. Nymphs aquatic, now populating almost exclusively running or cold water, often bearing external gills of different structure and disposition.

<==Plecoptera [Perlida, Perlina, Perlopseina]
    |--Ohionympha S02b [Palaeonemouridae GE05]
    |--Palaeoperlidae GE05
    |--Perlopsis S02a [Perlopseidae GE05]
    |    `--P. filicornis S02a
    `--+--Grypopterygomorpha GE05
       `--Arctoperlaria GE05
            |--Systellognatha [Perlomorpha] GE05
            |    |  i. s.: Sylvoperlodes [Tshekardoperlidae] S02a
            |    |           `--S. zhiltzova S02a
            |    |--Platyperla S02a [Platyperlidae GE05]
            |    |    `--P. platypoda S02a
            |    |--+--Pteronarcys TW05 [Pteronarcyidae GE05]
            |    |  |    `--P. californica TW05
            |    |  `--+--Cerconychia TW05 [Styloperlidae GE05]
            |    |     `--Tallaperla TW05 [Peltoperlidae GE05]
            |    |          `--T. lobata TW05
            |    `--+--Isoperla TW05 [Perlodidae GE05]
            |       |    |--I. davisi TW05
            |       |    |--I. grammatica B86
            |       |    `--I. luzoni Tierno de Figueroa & Vinçon 2005 F05
            |       `--+--Perlidae S02a
            |          |    |--Oyamia YS01
            |          |    |--Calineuria californica TW05
            |          |    `--Perla pictetii Lucas 1847 E12
            |          `--Chloroperlidae S02a
            |               |--Alloperla YS01
            |               `--Plumiperla diversa TW05
            `--Euholognatha [Nemourina] GE05
                 |  i. s.: Barathronympha victima S02a
                 |         Mesoleuctridae S02a
                 |           |--Mesoleuctra tibialis S02a
                 |           `--Mesoleuctroides saturatus S02a
                 |--Scopura TW05 [Scopuridae GE05]
                 |    `--S. montana TW05
                 `--+--Oemopteryx TW05 [Taeniopterygidae GE05]
                    |    `--O. vanduzeei TW05
                    |--Perlariopseidae GE05
                    |    |--Dicronemoura furcata S02a
                    |    |--Perlariopsis stipitata S02a
                    |    `--Spinoperla spinosa S02a
                    `--+--+--Aphanicerca TW05 [Notonemouridae GE05]
                       |  |    `--A. capensis TW05
                       |  `--Nemouridae GE05
                       |       |--Nemoura YS01
                       |       |--Malenka californica TW05
                       |       `--Protonemura zhiltzovae Vinçon & Ravizza 2005 F05
                       `--+--Baleyopteryx [Baleyopterygidae] S02a
                          |    `--B. orthoclada S02a
                          `--+--Capniidae GE05
                             |    |--Dobbertinopteryx S02a
                             |    |--Capnia gracilaria TW05
                             |    `--Capnura N88
                             `--Leuctridae GE05
                                  |--Leuctra major B86
                                  `--Paraleuctra vershina TW05

Plecoptera incertae sedis:
  Plutopteryx beata S02b
  Ecdyoperla S02a
  Gurvanopteryx Z02
  Triassoperla S02b
  Berekia S02b
  Perlitodes S02b
  Dipsoperla S02b
  Savina S02b
  Lycoleuctra S02b
  Uroperla S02b
  Flexoperla S02b
  Brachyptera tristis B86
  Paraperla frontalis B86
  Isocapnia B86
  Sharaperla S02c
  Bestioperlisca S02c
  Mongolonemoura S02c
  Megateptoperla grandis O81
  Perlisca S02b
  Perlomimus S02b
  Trianguliperla S02b
  Acroneuria californica R00

*Type species of generic name indicated


[B86] Botosaneanu, L. 1986. Insecta: Plecoptera, Trichoptera. In Stygofauna Mundi: A Faunistic, Distributional, and Ecological Synthesis of the World Fauna inhabiting Subterranean Waters (including the Marine Interstitial) (L. Botosaneanu, ed.) p. 619. E. J. Brill / Dr. W. Backhuys: Leiden.

[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.

[F05] Fernández, J. 2005. Noticia de nuevos táxones para la ciencia en el ámbito Íbero-Balear y Macaronésico. Nuevos táxones animales descritos en la península Ibérica y Macaronesia desde 1994 (IX). Graellsia 61 (2): 261-282.

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

[N88] Noonan, G. R. 1988. Biogeography of North American and Mexican insects, and a critique of vicariance biogeography. Systematic Zoology 37 (4): 366-384.

[O81] O'Brien, C. O. 1981. A. A. Book of New Zealand Wildlife: A guide to the native and introduced animals of New Zealand. Lansdowne Press: Auckland.

[R00] Ross, E. S. 2000. Embia: Contributions to the biosystematics of the insect order Embiidina. Occasional Papers of the California Academy of Sciences 149: 1-53, 1-36.

[S02a] Sinitshenkova, N. D. 2002a. Order Perlida Latreille, 1810. The stoneflies (=Plecoptera Burmeister, 1839). In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 281-287. Kluwer Academic Publishers: Dordrecht.

[S02b] Sinitshenkova, N. D. 2002b. 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.

[S02c] Sinitshenkova, N. D. 2002c. New late Mesozoic mayflies from the Shar-Teeg locality, Mongolia (Insecta, Ephemerida = Ephemeroptera). Paleontologicheskii Zhurnal 2002 (3): 43-48 (transl. Paleontological Journal 36 (3): 270-276).

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

[YS01] Yoshizawa, K., & T. Saigusa. 2001. Phylogenetic analysis of paraneopteran orders (Insecta: Neoptera) based on forewing base structure with comments on monophyly of Auchenorrhycha (Hemiptera). Systematic Entomology 26: 1-13.

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In History of Insects (A. P. Rasnitsyn & D. L. J. Quicke, eds) pp. 331-388. Kluwer Academic Publishers: Dordrecht.

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