Ensifera

Ensifera is a suborder of insects that includes the various types of crickets and their allies including: true crickets, camel crickets, bush crickets or katydids, grigs, weta and Cooloola monsters. This and the suborder Caelifera (grasshoppers and their allies) make up the order Orthoptera. Ensifera is believed to be a more ancient group than Caelifera, with its origins in the Carboniferous period,[2] the split having occurred at the end of the Permian period.[3] Unlike the Caelifera, the Ensifera contain numerous members that are partially carnivorous, feeding on other insects, as well as plants.

For the avian genus Ensifera, see Sword-billed hummingbird.

Ensifera
Temporal range: Artinskian/Kungurian–Holocene,
A bush cricket or katydid
Scientific classification Edit this classification
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Ensifera
Superfamilies and families

See text

Ensifer is Latin for "sword bearer", and refers to the typically elongated and blade-like ovipositor of the females.[4]

Characteristics

Characteristics shared by the two orthopteran suborders, Caelifera and Ensifera, are the mouthparts adapted for biting and chewing, the modified prothorax, the hind legs modified for jumping, the wing shape and venation, and the sound-producing stridulatory organs.[2]

Ensiferans are distinguished from Caeliferans by their elongated, threadlike antennae, which are often longer than the length of their bodies and have over 30 segments (except in the subterranean Cooloolidae family). For this reason, they are sometimes referred to as "long-horned orthopterans". In the families in which the males sing, the fore wings have modifications that include toothed veins and scrapers for making the noise, and the surrounding membranous areas amplify the sound. In these groups, the sound-detecting tympanal organs are located on the tibiae of the front legs.[5] The tarsi have three segments and the ovipositor is blade-like or needle-like. The male attaches the spermatophore externally to the female's gonopore. The spermatophore is often surrounded by a proteinaceous spermatophylax, the function of which is to provide a nutritional nuptial gift to the female.[5][6]

Taxonomy
A mole cricket, showing the front legs specialised for digging
A cave cricket, showing the long hind legs and antennae
A Cooloola monster, a subterranean family from Queensland, Australia
A splay-footed cricket from South India

The Orthoptera Species File database lists the following superfamilies and families.[7]

  • Infraorder †Elcanidea
    • Superfamily †Elcanoidea
      • Family †Elcanidae (Late Triassic - Paleocene)
      • Family †Permelcanidae (Early Permian - Late Triassic)
    • Superfamily †Permoraphidioidea
      • Family †Permoraphidiidae (Permian)
      • Family †Pseudelcanidae (Early Permian)
      • Family †Thueringoedischiidae (Early Permian)
      • Family incertae sedis
        • Genus †Acridiites Heer, 1865
  • Infraorder Gryllidea
  • Infraorder †Oedischiidea
    • Superfamily †Oedischioidea Handlirsch, 1906
      • Family †Anelcanidae Carpenter, 1986
      • Family †Bintoniellidae Handlirsch, 1939
      • Family †Mesoedischiidae Gorochov, 1987
      • Family †Oedischiidae Handlirsch, 1906
      • Family †Proparagryllacrididae Riek, 1956
      • Family †Pruvostitidae Zalessky, 1929
      • Family incertae sedis
        • Genus †Crinoedischia Béthoux & Beckemeyer, 2007
        • Genus †Loxoedischia Beckemeyer, 2011
    • Superfamily †Triassomantoidea Tillyard, 1922
      • Family †Adumbratomorphidae Gorochov, 1987
      • Family †Triassomantidae Tillyard, 1922
    • Superfamily †Xenopteroidea Riek, 1955
      • Family †Xenopteridae Riek, 1955
    • Superfamily incertae sedis
      • family †Permotettigoniidae Nel & Garrouste, 2016
      • Family incertae sedis
        • Genus †Permophyllum Prokop, et al, 2015
  • Infraorder Tettigoniidea
    • Superfamily Hagloidea - grigs
      • Family †Eospilopteronidae Cockerell, 1916
      • Family †Haglidae Handlirsch, 1906
      • Family †Hagloedischiidae Gorochov, 1986
      • Family †Prezottophlebiidae Martins-Neto, 2007
      • Family Prophalangopsidae Kirby, 1906
      • Family †Tuphellidae Gorochov, 1988
    • Superfamily †Phasmomimoidea Sharov, 1968
      • Family †Phasmomimidae Sharov, 1968
    • Superfamily Rhaphidophoroidea Walker, 1869
      • Family Rhaphidophoridae Walker, 1869 - camel crickets, cave crickets, cave wētā
    • Superfamily Schizodactyloidea Blanchard, 1845
    • Superfamily Stenopelmatoidea Burmeister, 1838
    • Superfamily Tettigonioidea Krauss, 1902
      • Family †Haglotettigoniidae Gorochov, 1988
      • Family Tettigoniidae - bush crickets, katydids, koringkrieks
    • Superfamily Incertae sedis
      • Family incertae sedis
        • Genus †Tettoraptor Gorochov, 2012
  • Infraorder incertae sedis
    • Superfamily †Gryllavoidea Gorochov, 1986
      • Family †Gryllavidae Gorochov, 1986
  • Superfamily Incertae sedis

Phylogeny
Prophalangopsidae: Jurassic fossil Pycnophlebia speciosa

The phylogenetic relationships of the Ensifera, summarized by Darryl Gwynne in 1995 from his own work and that of earlier authors,[lower-alpha 1] are shown in the following cladogram, with the Orthoptera divided into two main groups, Ensifera and Caelifera (grasshoppers). Fossil Ensifera are found from the late Carboniferous period onwards.[5][10]

The oldest known fossil in the Archaeorthoptera, the crown group of the Orthoptera, and also the oldest member of the Pterygota (winged insects), is from the Namurian (324 mya) Lower Carboniferous beds in the Upper Silesian Basin of the Czech Republic.[11]

Orthoptera
Ensifera
Elcanidea

†Elcanoidea

†Permoraphidioidea

Oedischiidea

†Oedischioidea

†Triassomantoidea

†Xenopteroidea

Tettigoniidea

Hagloidea: (including grigs)

†Phasmomimoidea

Stenopelmatoidea (wētā, king crickets)

Tettigonioidea (bush crickets, katydids, koringkreiks)

Rhaphidophoroidea (cave wētā, cave crickets)

†Gryllavoidea

Grylloidea (crickets)

Schizodactyloidea (dune crickets)

Caelifera

(grasshoppers, groundhoppers, pygmy mole crickets)

Notes
  1. Gwynne cites Ander 1939, Zeuner 1939, Judd 1947, Key 1970, Ragge 1977 and Rentz 1991 as supporting the two-part scheme (Ensifera, Caelifera) in his 1995 paper.[10]

References
  1. Wang, Yan-hui; Engel, Michael S.; Rafael, José A.; Wu, Hao-yang; Rédei, Dávid; Xie, Qiang; Wang, Gang; Liu, Xiao-guang; Bu, Wen-jun (2016). "Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda)". Scientific Reports. 6: 38939. doi:10.1038/srep38939. PMC 5154178. PMID 27958352.
  2. Resh, Vincent H.; Cardé, Ring T. (2009). Encyclopedia of Insects. Academic Press. pp. 232, 733–735. ISBN 978-0-08-092090-0.
  3. Zeuner, F. E. (1939). Fossil Orthoptera Ensifera. London: British Museum Natural History.
  4. "Orthoptera". Natural History Museum. Retrieved 2015-05-13.
  5. Gwynne, Darryl T.; DeSutter; Laure (1996). "Ensifera: Crickets, katydids and weta". TOLweb. Retrieved 8 May 2015.
  6. Vahed; K. (1998). "The function of nuptial feeding in insects: review of empirical studies" (PDF). Biological Reviews. 73: 43–78. doi:10.1111/j.1469-185X.1997.tb00025.x. S2CID 86644963. Archived from the original (PDF) on 2012-03-11. Retrieved 2015-05-08.
  7. "Suborder Ensifera". Orthoptera Species File. Retrieved 9 May 2015.
  8. Orthoptera Species File (Version 5.0/5.0)
  9. Archibald, S. B.; Gu, J.-J.; Mathewes, R. W. (2022). "The Palaeorehniidae (Orthoptera, Ensifera, "Zeuneropterinae"), and new taxa from the early Eocene Okanagan Highlands, western North America". Zootaxa. 5100 (4): 559–572. doi:10.11646/zootaxa.5100.4.6.
  10. Gwynne, Darryl T. (1995). "Phylogeny of the Ensifera (Orthoptera): a hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care in crickets, katydids, and weta". Journal of Orthoptera Research. 4 (4): 203–218. doi:10.2307/3503478. JSTOR 3503478.
  11. Prokop, Jakub; Nel, André; Hoch, Ivan (2005). "Discovery of the oldest known Pterygota in the Lower Carboniferous of the Upper Silesian Basin in the Czech Republic (Insecta: Archaeorthoptera)". Geobios. 38 (3): 383–387. doi:10.1016/j.geobios.2003.11.006.
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