Caelifera

The Caelifera are a suborder of orthopteran insects.[2] They include the grasshoppers and grasshopper-like insects, as well as other superfamilies classified with them: the ground-hoppers (Tetrigoidea) and pygmy mole crickets (Tridactyloidea). The latter should not be confused with the mole crickets (Gryllotalpidae), which belong to the other Orthopteran sub-order Ensifera.

Caelifera
Temporal range: Changhsingian–Present
A pygmy mole-cricket
A groundhopper
Scientific classification
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Orthoptera
Suborder: Caelifera
Ander, 1939
Infraorders[1]

Subdivisions and their distribution

The Caelifera include some 2,400 valid genera and about 12,000 known species. Many undescribed species probably exist, especially in tropical forests. The Caelifera have a predominantly tropical distribution (as with most Orthoptera) with fewer species known from temperate climate zones. Caelifera are divided into two infraorders: the more basal Tridactylidea and the Acrididea or grasshopper-like species. This latter name is derived from older sources, such as Imms,[3] which placed the "short-horned grasshoppers" and locusts at the family level (Acrididae).

Affiliations

The phylogeny of the Caelifera, is described in detail for grasshoppers, with 6 out of 8 extant superfamilies shown here as a cladogram. Like the Ensifera, Caelifera and all of its superfamilies appear to be monophyletic.[2][7]

Orthoptera
Ensifera (crickets, etc.)

[6 superfamilies]

Caelifera
 Tridactylidea 

Tridactyloidea

Dzhajloutshelloidea (extinct)

 Acrididea 

Tetrigoidea

 Acridomorpha 

Eumastacoidea

Pneumoroidea

Pyrgomorphoidea

Acridoidea etc.

The phylogeny of the Caelifera, based on mitochondrial ribosomal RNA of thirty-two taxa in six out of seven superfamilies, is shown as a cladogram. The Ensifera, Caelifera and all the superfamilies of grasshoppers except Pamphagoidea appear to be monophyletic.[8][9]

Orthoptera
Ensifera (crickets)

[6 superfamilies]

Caelifera

Tridactyloidea

Tetrigoidea

Eumastacidae

Proscopiidae

Pneumoridae

Pyrgomorphidae

Acrididae + Pamphagidae

Fossil grasshoppers at the Royal Ontario Museum

In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the Permo-Triassic boundary;[10] the earliest insects that are certainly Caeliferans are Eolocustopsis from the latest Permian (Changhsingian) of the Beaufort Group, South Africa[11] and Locustavidae from the early Triassic, roughly 250 million years ago. The oldest known member of Caelifera is The group diversified during the Triassic and have remained important plant-eaters from that time to now. The first modern families such as the Eumastacidae, Tetrigidae and Tridactylidae appeared in the Cretaceous, though some insects that might belong to the last two of these groups are found in the early Jurassic.[12][13] Morphological classification is difficult because many taxa have converged towards a common habitat type; recent taxonomists have concentrated on the internal genitalia, especially those of the male. This information is not available from fossil specimens, and the palaentological taxonomy is founded principally on the venation of the hindwings.[2]

The Caelifera includes some 2,400 valid genera and about 11,000 known species. Many undescribed species probably exist, especially in tropical wet forests. The Caelifera have a predominantly tropical distribution with fewer species known from temperate zones, but most of the superfamilies have representatives worldwide. They are almost exclusively herbivorous and are probably the oldest living group of chewing herbivorous insects.[2]

The most diverse superfamily is the Acridoidea, with around 8,000 species. The two main families in this are the Acrididae (grasshoppers and locusts) with a worldwide distribution, and the Romaleidae (lubber grasshoppers), found chiefly in the New World. The Ommexechidae and Tristiridae are South American, and the Lentulidae, Lithidiidae and Pamphagidae are mainly African. The Pauliniids are nocturnal and can swim or skate on water, and the Lentulids are wingless.[12] Pneumoridae are native to Africa, particularly southern Africa, and are distinguished by the inflated abdomens of the males.[14]

Economic significance and terminology

A number of species, especially in the Acridoidea, are significant agricultural pests, but not all of them are locusts: a non-taxonomic term referring to species whose populations which may change morphologically when crowded and show swarming behaviour.[15] Examples of agricultural grasshopper pests that are not called locusts include the Senegalese grasshopper and certain species in the Pyrgomorphidae, notably the variegated grasshopper (Zonocerus variegatus).

References

  1. ITIS: Caelifera (retrieved 2 August 2017)
  2. Rowell, Hugh; Flook, Paul (2001). "Caelifera: Shorthorned Grasshoppers, Locusts and Relatives". Tree of Life web project. Archived from the original on 8 April 2015. Retrieved 3 April 2015.
  3. Imms AD, rev. Richards OW & Davies RG (1970) A General Textbook of Entomology 9th Ed. Methuen 886 pp.
  4. Ragge DR (1965). Grasshoppers, Crickets & Cockroaches of the British Isles. F Warne & Co, London. p. 299.
  5. ITIS: Acrididea (Retrieved 23/7/2017)
  6. Orthoptera Species File: infraorder Acrididea (Retrieved 20/7/2017)
  7. Flook, P. K.; Rowell, C. H. F. (1997). "The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences". Molecular Phylogenetics and Evolution. 8 (1): 89–103. doi:10.1006/mpev.1997.0412. PMID 9242597.
  8. Flook, P.K.; Rowell, C.H.F. (1997). "The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences". Molecular Phylogenetics and Evolution. 8 (1): 89–103. doi:10.1006/mpev.1997.0412. PMID 9242597.
  9. Zhang, Hong-Li; Huang, Yuan; Lin, Li-Liang; Wang, Xiao-Yang; Zheng, Zhe-Min (2013). "The phylogeny of the Orthoptera (Insecta) as deduced from mitogenomic gene sequences". Zoological Studies. 52: 37. doi:10.1186/1810-522X-52-37.
  10. Zeuner, F.E. (1939). Fossil Orthoptera Ensifera. British Museum Natural History. OCLC 1514958.
  11. Song, Hojun; Amédégnato, Christiane; Cigliano, Maria Marta; Desutter-Grandcolas, Laure; Heads, Sam W.; Huang, Yuan; Otte, Daniel; Whiting, Michael F. (December 2015). "300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling". Cladistics. 31 (6): 621–651. doi:10.1111/cla.12116.
  12. Grimaldi, David; Engel, Michael S. (2005). Evolution of the Insects. Cambridge University Press. p. 210. ISBN 978-0-521-82149-0. Archived from the original on 27 November 2017.
  13. Béthoux, Oliver; Ross, A.J. (2005). "Mesacridites Riek, 1954 (Middle Triassic; Australia) transferred from Protorthoptera to Orthoptera: Locustavidae". Journal of Paleontology. 79 (3): 607–610. doi:10.1666/0022-3360(2005)079<0607:mrmatf>2.0.co;2.
  14. Donelson, Nathan C.; van Staaden, Moira J. (2005). "Alternate tactics in male bladder grasshoppers Bullacris membracioides (Orthoptera: Pneumoridae)" (PDF). Behaviour. 142 (6): 761–778. doi:10.1163/1568539054729088. Archived from the original (PDF) on 20 December 2016.
  15. Uvarov BP (1966) Grasshoppers & Locusts. A Handbook of General Acridology Cambridge University Press, London 1:481 pp.
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