caelifera

{{Short description|Suborder of insects}}

{{About|the orthopteran suborder|the blowfly genus with a similar name|Calliphora}}

{{Automatic taxobox

| fossil_range = {{fossilrange|Changhsingian|Present}}

| image = Pygmy mole cricket (8071068977) cropped.jpg

| image_caption = A pygmy mole-cricket

| image2 = Melanoplus femurrubrum 20070903.jpg

| image2_caption = Red-legged grasshopper

| taxon = Caelifera

| authority = Ander, 1939

| subdivision_ranks = Infraorders

| subdivision_ref = [https://www.itis.gov/servlet/SingleRpt/SingleRpt?search_topic=TSN&search_value=102161#null ITIS: Caelifera (retrieved 2 August 2017)]

| subdivision = * Acrididea

}}

The Caelifera are a suborder of orthopteran insects. 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.

The name of this suborder comes from Latin meaning chisel-bearing ("chisel" in Latin: caelum),{{cite encyclopedia |url=https://www.perseus.tufts.edu/hopper/text?doc=Perseus:text:1999.04.0059:entry=caelum1&highlight=chisel |title= caelum |last1= Lewis |first1= Charlton T. |last2= Short |first2= Charles |dictionary= A Latin Dictionary |edition= |publisher= Perseus Digital Library |date = 1879 }} referring to the "stout" shape of its species' ovipositors.{{cite book| last=Himmelman| first=John| title=Cricket Radio| url=https://books.google.com/books?id=qp7Q4Vn9l-oC&pg=PA45| year=2011| publisher=Harvard University Press| isbn=978-0-674-06102-6| page=45| url-status=live| archive-url=https://web.archive.org/web/20171127023307/https://books.google.com/books?id=qp7Q4Vn9l-oC&pg=PA45| archive-date=27 November 2017}}

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,Imms AD, rev. Richards OW & Davies RG (1970) A General Textbook of Entomology 9th Ed. Methuen 886 pp. 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 six out of eight extant superfamilies shown here as a cladogram. Like the Ensifera, Caelifera and all of its superfamilies appear to be monophyletic.{{cite journal |last1=Flook |first1=P. K. |last2=Rowell |first2=C. H. F. |title=The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences |journal=Molecular Phylogenetics and Evolution |date=1997 |volume=8 |issue=1 |pages=89–103 |doi=10.1006/mpev.1997.0412 |pmid=9242597|bibcode=1997MolPE...8...89F }}

{{clade| style=font-size:90%;line-height:90%

|label1=Orthoptera

|1={{clade

|label1=Ensifera (crickets, etc.)

|1=[6 superfamilies] 70px

|label2=Caelifera

|2={{clade

|label1= Tridactylidea

|1=Tridactyloidea 70px

|label2= Acrididea

|2={{clade

|1=Tetrigoidea 70px

|label3= Acridomorpha

|3={{clade

|1=Eumastacoidea 70px

|2={{clade

|1=Pneumoroidea 70px

|2={{clade

|1=Pyrgomorphoidea 70px

|2=Acridoidea etc. 70px

}}

}}

}}

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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.{{cite journal |last1=Flook |first1=P.K. |last2=Rowell |first2=C.H.F. |title=The Phylogeny of the Caelifera (Insecta, Orthoptera) as Deduced from mtrRNA Gene Sequences |journal=Molecular Phylogenetics and Evolution |date=1997 |volume=8 |issue=1 |pages=89–103 |doi=10.1006/mpev.1997.0412 |pmid=9242597|bibcode=1997MolPE...8...89F }}{{cite journal |last1=Zhang |first1=Hong-Li|last2=Huang |first2=Yuan|last3=Lin |first3=Li-Liang |last4=Wang |first4=Xiao-Yang |last5=Zheng|first5=Zhe-Min |title=The phylogeny of the Orthoptera (Insecta) as deduced from mitogenomic gene sequences |journal=Zoological Studies |date=2013 |volume=52 |page=37 |doi=10.1186/1810-522X-52-37|doi-access=free }}

{{clade| style=font-size:85%;line-height:85%

|label1=Orthoptera

|1={{clade

|label1=Ensifera (crickets)

|1=[6 superfamilies] 35px

|label2=Caelifera

|2={{clade

|1=Tridactyloidea 50px

|2={{clade

|1=Tetrigoidea 50px

|2={{clade

|1=Eumastacidae 80px

|2={{clade

|1=Proscopiidae 50px

|2={{clade

|1=Pneumoridae 50px

|2={{clade

|1=Pyrgomorphidae 50px

|2=Acrididae + Pamphagidae 50px

}}

}}

}}

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File:Fossil grasshoppers - Royal Ontario Museum - DSC00013.JPG]]

In evolutionary terms, the split between the Caelifera and the Ensifera is no more recent than the Permo-Triassic boundary;{{cite book |author=Zeuner, F.E. |date=1939 |title=Fossil Orthoptera Ensifera |publisher=British Museum Natural History |oclc=1514958}} the earliest insects that are certainly Caeliferans are Eolocustopsis of family Eolocustopsidae from the latest Permian (Changhsingian) of the Beaufort Group, South Africa,{{Cite journal|last1=Song|first1=Hojun|last2=Amédégnato|first2=Christiane|last3=Cigliano|first3=Maria Marta|last4=Desutter-Grandcolas|first4=Laure|last5=Heads|first5=Sam W.|last6=Huang|first6=Yuan|last7=Otte|first7=Daniel|last8=Whiting|first8=Michael F.|date=December 2015|title=300 million years of diversification: elucidating the patterns of orthopteran evolution based on comprehensive taxon and gene sampling|journal=Cladistics|language=en|volume=31|issue=6|pages=621–651|doi=10.1111/cla.12116|pmid=34753270 |s2cid=53702892 |doi-access=free|hdl=11336/10677|hdl-access=free}} followed by Locustavidae and Dzhajloutshellidae from the mid-Triassic (Ladinian age), roughly 242 to 237 million years ago.{{cite journal|last1=Xu|first1=Chunpeng|last2=Fang|first2=Yan|last3=Heads|first3=Sam|last4=Zhang|first4=Qiang|last5=Jarzembowski|first5=Edmund A.|last6=Wang|first6=Bo|title=The first Orthoptera (Insecta) from the Triassic of China|journal=Alcheringa|volume=44|year=2020|issue=1|pages=93-98|doi=10.1080/03115518.2019.1662485}}{{cite journal|last=Shcherbakov|first=Dmitry E.|title=Madygen, Triassic Lagerstätte number one, before and after Sharov|journal=Alavesia|volume=2|pages=113-124|year=2008|s2cid=55131160}} 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.{{cite book |author1=Grimaldi, David |author2=Engel, Michael S. |title=Evolution of the Insects |url=https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA210 |year=2005 |publisher=Cambridge University Press |isbn=978-0-521-82149-0 |page=210 |url-status=live |archive-url=https://web.archive.org/web/20171127023307/https://books.google.com/books?id=Ql6Jl6wKb88C&pg=PA210 |archive-date=27 November 2017 }}{{cite journal |last1=Béthoux |first1=Oliver |last2=Ross |first2=A.J. |title=Mesacridites Riek, 1954 (Middle Triassic; Australia) transferred from Protorthoptera to Orthoptera: Locustavidae |journal=Journal of Paleontology |date=2005|volume=79 |issue=3 |pages=607–610 |doi=10.1666/0022-3360(2005)079<0607:mrmatf>2.0.co;2|bibcode=2005JPal...79..607B |s2cid=131591210 }} 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 paleontological taxonomy is founded principally on the venation of the hindwings.

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.{{cite web |url=http://tolweb.org/tree?group=Caelifera |title=Caelifera: Shorthorned Grasshoppers, Locusts and Relatives |author1=Rowell, Hugh |author2=Flook, Paul |year=2001 |work=Tree of Life web project |access-date=3 April 2015 |url-status=live |archive-url=https://web.archive.org/web/20150408064407/http://tolweb.org/tree?group=Caelifera |archive-date=8 April 2015 }}

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. Pneumoridae are native to Africa, particularly southern Africa, and are distinguished by the inflated abdomens of the males.{{cite journal |last1=Donelson |first1=Nathan C. |last2=van Staaden |first2=Moira J. |year=2005 |title=Alternate tactics in male bladder grasshoppers Bullacris membracioides (Orthoptera: Pneumoridae) |journal=Behaviour |volume=142 |issue=6 |pages=761–778 |doi=10.1163/1568539054729088 |url=http://caspar.bgsu.edu/~mooi/Home/Publications_files/MvS_Behav05.pdf |url-status=dead |archive-url=https://web.archive.org/web/20161220162119/http://caspar.bgsu.edu/~mooi/Home/Publications_files/MvS_Behav05.pdf |archive-date=20 December 2016 }}

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.Uvarov BP (1966) Grasshoppers & Locusts. A Handbook of General Acridology Cambridge University Press, London 1:481 pp. 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).

See also

References

{{Reflist}}