Editing Orthoptera

{{short description|Order of insects that includes grasshoppers}}
{{Automatic taxobox
| taxon = Orthoptera
| fossil_range= [[Carboniferous]]–recent {{fossilrange|359|0}}
| image = Roesel's bush-cricket (Metrioptera roeselii diluta) male.jpg
| image_caption = [[Roesel's bush-cricket]]<br />family [[Tettigoniidae]]
| authority = [[Latreille]], 1793
| display_parents = 2
| subdivision_ranks = Extant [[suborder]]s and [[Taxonomic rank|superfamilies]]
| subdivision = 
Suborder [[Ensifera]]
*[[Grylloidea]]
*[[Gryllotalpoidea]]
*[[Hagloidea]]
*[[Rhaphidophoroidea]]
*[[Schizodactyloidea]]
*[[Stenopelmatoidea]]
*[[Tettigonioidea]]
Suborder [[Caelifera]]
*[[Acridoidea]]
*[[Eumastacoidea]]
*[[Pneumoroidea]]
*[[Pyrgomorphoidea]]
*[[Tanaoceroidea]]
*[[Tetrigoidea]]
----
*[[Tridactyloidea]]
*[[Trigonopterygoidea]]
}}

'''Orthoptera''' ({{etymology|grc|''{{wikt-lang|grc|ὀρθός}}'' ({{grc-transl|ὀρθός}})|straight||''{{wikt-lang|grc|πτερά}}'' ({{grc-transl|πτερά}})|wings}}) is an [[order (biology)|order]] of [[insect]]s that comprises the [[grasshopper]]s, [[locust]]s, and [[cricket (insect)|cricket]]s, including closely related insects, such as the [[Tettigoniidae|bush crickets or katydids]] and [[wētā]]. The order is subdivided into two suborders: [[Caelifera]] – grasshoppers, locusts, and close relatives; and [[Ensifera]] – crickets and close relatives.

More than 20,000 [[species]] are distributed worldwide.<ref>{{Cite web|title = Orthoptera - Grasshoppers, Locusts, Crickets, Katydids |url= http://www.discoverlife.org/mp/20q?search=Orthoptera |website = Discover Life |access-date = 2017-09-06}}</ref> The insects in the order have [[incomplete metamorphosis]], and produce [[sound]] (known as a "[[stridulation]]") by rubbing their wings against each other or their legs, the wings or legs containing rows of corrugated bumps. The [[Tympanum (anatomy)|tympanum]], or [[ear]], is located in the front [[Tibia (arthropod)|tibia]] in crickets, [[mole cricket]]s, and bush crickets or katydids, and on the first abdominal segment in the grasshoppers and locusts.<ref name=IIBD/> These organisms use vibrations to locate other individuals.

Grasshoppers and other orthopterans are able to fold their [[Insect wing|wings]] (i.e. they are members of [[Neoptera]]).

==Etymology==
The name is derived from the [[Ancient Greek|Greek]] {{Lang|grc|ὀρθός}} {{Transliteration|grc|orthos}} meaning "straight" and {{Lang|grc|πτερόν}} ''{{Transliteration|grc|pteron}}'' meaning "wing".

==Characteristics==
Orthopterans have a generally [[cylindrical]] body, with elongated hindlegs and musculature adapted for [[jumping]]. They have [[mandible (arthropod)|mandibulate]] mouthparts for biting and chewing and large [[compound eye]]s, and may or may not have [[ocelli]], depending on the species. The [[antenna (biology)|antennae]] have multiple joints and filiform type, and are of variable length.<ref name=IIBD>{{cite book |author= Hoell, H.V., Doyen, J.T. & Purcell, A.H. |year=1998 |title=Introduction to Insect Biology and Diversity, 2nd ed. |publisher= Oxford University Press |pages= 392–394|isbn= 978-0-19-510033-4}}</ref>

The first and third segments on the [[Thorax (insect anatomy)|thorax]] are larger, while the second segment is much smaller. They have two pairs of [[insect wing|wings]], which are held overlapping the [[abdomen]] at rest. The forewings, or [[tegmina]], are narrower than the hindwings and hardened at the base, while the hindwings are membranous, with straight veins and numerous cross-veins. At rest, the hindwings are held folded fan-like under the forewings. The final two to three segments of the abdomen are reduced, and have single-segmented [[Cercus|cerci]].<ref name=IIBD/>

==Life cycle==
Orthopterans have a paurometabolous lifecycle or [[incomplete metamorphosis]]. The use of sound is generally crucial in courtship, and most species have distinct songs.<ref>{{citation |url=https://books.google.com/books?id=SHHQVehP488C&pg=PA74 |title=The practical entomologist |pages=74–75 |first=Rick |last=Imes |publisher=Simon and Schuster |isbn=978-0-671-74695-7 |year= 1992 }}</ref> Most grasshoppers lay their [[egg (biology)|eggs]] in the ground or on vegetation. The eggs hatch and the young [[Nymph (biology)|nymphs]] resemble adults, but lack wings and at this stage are often called 'hoppers'. They may often also have a radically different coloration from the adults. Through successive [[ecdysis|moults]], the nymphs develop wings until their final moult into a mature adult with fully developed wings.<ref name=IIBD/>

The number of moults varies between species; growth is also very variable and may take a few weeks to some months depending on food availability and weather conditions.

==Evolution==

This order evolved {{Ma|300}} with a division into two suborders – [[Caelifera]] and [[Ensifera]] – occurring {{Ma|256}}.<ref name=Chang2020>Chang H, Qiu Z, Yuan H, Wang X, Li X, Sun H, Guo X, Lu Y, Feng X, Majid M, Huang Y (2020) Evolutionary rates of and selective constraints on the mitochondrial genomes of Orthoptera insects with different wing types. Mol Phylogenet Evol </ref>

===Phylogeny===

The Orthoptera are divided into two suborders, [[Caelifera]] and [[Ensifera]], that have been shown to be [[monophyletic]].<ref name="Zhou2010">Zhou Z, Ye H, Huang Y, Shi F. (2010) The phylogeny of Orthoptera inferred from mtDNA and description of ''Elimaea cheni'' (Tettigoniidae: Phaneropterinae) mitogenome. ''J. Genet. Genomics.'' 37(5):315-324</ref><ref name=Gwynne>{{cite journal |last1=Gwynne |first1=Darryl T. |title=Phylogeny of the Ensifera (Orthoptera): a hypothesis supporting multiple origins of acoustical signalling, complex spermatophores and maternal care in crickets, katydids, and weta |journal= Journal of Orthoptera Research|date=1995 |volume=4 |issue=4 |pages=203–218 |doi=10.2307/3503478 |jstor=3503478}}</ref><ref>{{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 }}</ref> A recent comprehensive phylogeny based on analyses of data from transcriptomes and mitochondrial genomes found the following relationships within Orthoptera.<ref name=Song-2020>{{cite journal |last1=Song |first1=Hojun |last2=Béthoux |first2=Olivier |last3=Shin |first3=Seunggwan  |first4=Alexander |last4=Donath |first5=Harald |last5=Letsch |first6=Shanlin |last6=Liu |first7=Duane D. |last7=McKenna |first8=Guanliang |last8=Meng |first9=Bernhard |last9=Misof |first10=Lars |last10=Podsiadlowski |first11=Xin |last11=Zhou |first12=Benjamin |last12=Wipfler |first13=Sabrina |last13=Simon  |title=Phylogenomic analysis sheds light on the evolutionary pathways towards acoustic communication in Orthoptera. |journal=Nat. Commun. |volume=11 |pages=4939 |year=2020 |doi=10.1038/s41467-020-18739-4 |pmc=7532154 }}</ref>

{{clade| style=font-size:90%;line-height:90%
|label1='''Orthoptera'''
|1={{clade
   |label1=[[Ensifera]]
   |1={{clade
      |label1=[[Gryllidea]]
      |1={{clade |state1=double
         |1=[[Gryllotalpoidea]]{{efn|Gryllotalpoidea is paraphyletic with respect to Grylloidea, with either [[Gryllotalpidae]] or [[Myrmecophilidae]] as sister to Grylloidea<ref name=Song-2020/> }} (mole crickets and ant crickets)
         |2=[[Grylloidea]] ("true crickets", scaly crickets, wood crickets, etc) [[File:Arachnocephalus vestitus01.jpg|70px]]
         }}
      |label2=[[Tettigoniidea]]
      |2={{clade
         |1=[[Rhaphidophoroidea]] (cave crickets, camel crickets, spider crickets and sand treaders) [[File:Ceuthophiluscricket.jpg|70px]]
         |2={{clade  
            |1=[[Schizodactyloidea]] (dune crickets) [[File:Пальцепалый кузнечик.jpg|50px]]
            |2={{clade
               |1=[[Stenopelmatoidea]] (Jerusalem crickets, king crickets, leaf-rolling crickets and Cooloola monsters)
               |2={{clade
                  |1=[[Hagloidea]] (grigs)
                  |2=[[Tettigonioidea]] (katydids or bush crickets) [[File:Cricket September 2010-1.jpg|70px]] 
                  }}
               }}
            }}
         }}
      }}
   |label2=[[Caelifera]]
   |2={{clade
      |label1=[[Tridactylidea]]
      |1=[[Tridactyloidea]] [[File:Pygmy mole cricket (8071068977) cropped.jpg|70px]]
      |label2=[[Acrididea]]
      |2={{clade
         |1=[[Tetrigoidea]] [[File:Tetrix subulata 2.JPG|70px]]
         |label2= [[Acridomorpha]] |sublabel2=([[grasshopper]]s)
         |2={{clade
                |1={{clade |state1=double
                   |1=[[Eumastacoidea]]{{efn|Eumastacoidea is paraphyletic with respect to Proscopioidea, with the latter in a derived position as sister to [[Episactidae]].<ref name=Song-2020/> }} [[File: Monkey hopper (14795010039).jpg|70px]] 
                   |2=[[Proscopioidea]] 
                   }}
                |2={{clade
                   |1=[[Tanaoceroidea]]
                   |2={{clade
                      |1={{clade |state1=double
                         |1=[[Trigonopterygoidea]]{{efn|Trigonopterygoidea is paraphyletic with respect to Pneumoroidea, with [[Trigonopterygidae ]] as sister to [[Pneumoridae]].<ref name=Song-2020/>}}
                         |2=[[Pneumoroidea]] [[File:Bladder Grasshopper (Bullacris intermedia) (30068047440).jpg |70px]]
                         }}
                      |2={{clade
                         |1=[[Pyrgomorphoidea]] [[File:Variegated grasshopper (Zonocerus variegatus).jpg|70px]] 
                         |2=[[Acridoidea]] [[File:SGR laying.jpg|70px]]
                         }}
                      }}
                   }}
              }}
          }} 
       }}
   }}
}}

;Notes
{{notelist}}

===Taxonomy===
[[File:Garden locust (Acanthacris ruficornis).jpg|thumb|Garden locust (''Acanthacris ruficornis''), [[Ghana]], family [[Acrididae]]]]
[[File:Variegated grasshopper (Zonocerus variegatus).jpg|thumb|Variegated grasshopper (''Zonocerus variegatus''), [[Ghana]], family [[Pyrgomorphidae]]]]
[[File:Peruvian Stick Bug.jpg|thumb|[[Proscopiidae|Proscopiidae gen. sp.]] from the [[Andes]] of [[Peru]]]]

Taxonomists classify members of the Caelifera and Ensifera into infraorders and superfamilies as follows:<ref>{{cite web|title=Orthoptera Species File Online|url=https://bugscope.beckman.illinois.edu/pdfs/insects/Orthoptera_Species.pdf |publisher=University of Illinois|access-date=6 January 2018}}</ref><ref name="BlackithBlackith1968">{{cite journal|last1=Blackith|first1= RE|last2=Blackith|first2= RM|title=A numerical taxonomy of Orthopteroid insects|journal=Australian Journal of Zoology|volume=16|issue=1 |year=1968|pages=111 |doi=10.1071/ZO9680111}}</ref><ref name="FlookKlee1999">{{cite journal|last1=Flook|first1=P. K.|last2=Klee|first2=S. |last3=Rowell|first3=C. H. F.|last4=Simon|first4=C.|title=Combined Molecular Phylogenetic Analysis of the Orthoptera (Arthropoda, Insecta) and Implications for Their Higher Systematics|journal=Systematic Biology|volume=48 |issue=2|year=1999 |pages=233–253|issn=1076-836X|doi=10.1080/106351599260274|pmid=12066707|url=http://doc.rero.ch/record/300023/files/48-2-233.pdf|doi-access=free}}</ref><ref>{{Cite web |title=superfamily Gryllotalpoidea Leach, 1815: Orthoptera Species File |url=http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1128272 |access-date=2023-09-01 |website=orthoptera.speciesfile.org}}</ref>  

* Suborder [[Caelifera]] – grasshoppers, pygmy mole crickets and allies
** Infraorder [[Acrididea]]
*** Superfamily [[Acridoidea]] – grasshoppers, locusts
*** Superfamily [[Eumastacoidea]] – monkey or matchstick grasshoppers and allies
*** Superfamily [[Locustopsoidea]]†
*** Superfamily [[Pneumoroidea]] – bladder grasshoppers
*** Superfamily [[Pyrgomorphoidea]] – gaudy grasshoppers
*** Superfamily [[Tanaoceroidea]] – desert long-horned grasshoppers
*** Superfamily [[Tetrigoidea]] – ground-hoppers or grouse locusts
*** Superfamily [[Trigonopterygoidea]] – leaf grasshoppers
** Infraorder [[Tridactylidea]]
*** Superfamily [[Dzhajloutshelloidea]]†
*** Superfamily [[Regiatoidea]]†
*** Superfamily [[Tridactyloidea]] – pygmy mole crickets and allies
* Suborder [[Ensifera]] – crickets
** Superfamily [[Grylloidea]] – crickets 
** Superfamily [[Gryllotalpoidea]] – mole crickets and ant crickets
** Superfamily [[Hagloidea]] – grigs and allies
** Superfamily [[Phasmomimoidea]]†
** Superfamily [[Rhaphidophoroidea]] – camel crickets, cave crickets, cave wētā
** Superfamily [[Schizodactyloidea]] – dune crickets
** Superfamily [[Stenopelmatoidea]] – wētā and allies
** Superfamily [[Tettigonioidea]] – katydids / bush crickets
*''Incertae sedis''
**Superfamily [[Elcanidae|Elcanoidea]]† [[Permian]]-[[Paleocene]]

== Relationships with humans ==

===As pests===

Several species of Orthoptera are considered pests of crops and rangelands or seeking warmth in homes by humans. The two groups of Orthoptera that cause the most damage are [[grasshopper]]s and [[locust]]s. Locust are historically known for wiping out fields of crops in a day. Locust have the ability to eat up to their own body weight in a single day.<ref name=":0">{{Cite web |url=http://animals.nationalgeographic.com/animals/bugs/locust/ |archive-url=https://web.archive.org/web/20100207174802/http://animals.nationalgeographic.com/animals/bugs/locust/ |url-status=dead |archive-date=February 7, 2010 |title=Locusts, Locust Pictures, Locust Facts - National Geographic |last=Society |first=National Geographic |website=National Geographic |access-date=2016-04-11}}</ref> Individuals gather in large groups called swarms, these swarms can range up to 80 million individuals that stretch 460 square miles.<ref name=":0" /> Grasshoppers can cause major agricultural damage but not to the documented extent as locust historically have. These insects mainly feed on weeds and grasses, however, during times of drought and high population density they will feed on crops. They are a known pest in [[soybean]] fields and will likely feed on these crops once preferred food sources have become scarce.<ref>{{Cite web |url=https://extension.entm.purdue.edu/fieldcropsipm/insects/soybean-grasshopper.php |title=Grasshoppers {{!}} Pests {{!}} Soybean {{!}} Integrated Pest Management {{!}} IPM Field Crops {{!}} Purdue University |last=Krupke |first=Christian  |website=extension.entm.purdue.edu |access-date=2016-04-11}}</ref>

===As food===
{{See also|Insects as food}}
Most orthopterans are edible, making up 13% of all insects including some 80 species of grasshoppers being regularly consumed worldwide.<ref name=":1">{{Cite book |last=van Huis |first=Arnold |url=https://www.fao.org/3/i3253e/i3253e.pdf |title=Edible Insects: Future Prospects for Food and Feed Security |isbn=9789251075968 |location=Rome |oclc=868923724 |author-link=Arnold van Huis |pages=13–14}}</ref> In [[Madagascar]] and [[Oaxaca]], grasshoppers and locusts are usually collected early in the morning when it is cooler as the orthopterans are less mobile due to being [[Ectotherm|cold-blooded]].<ref name=":1" /> In Thailand, [[House cricket|house crickets]] are commonly reared and eaten; as of 2012, around 20,000 cricket farmers had farms in 53 of their [[Provinces of Thailand|76 provinces]].<ref name=":1" />

In the second century BCE in [[Ancient Greece]], [[Diodorus Siculus]] is known to have called people from [[Ethiopia]] ''Acridophagi'', meaning "eaters of locusts."<ref name=":1" />

In [[Judaism]], the Orthoptera include the only insects considered [[kosher]]. The list of dietary laws in the [[book of Leviticus]] forbids all flying insects that walk, but makes an exception for [[Kosher locust|certain locusts]].<ref name="Gordon">{{citation |title=The eat-a-bug cookbook |pages=3 |url=https://books.google.com/books?id=sI0Zax6ljq8C&pg=PA3 |first=David George |last=Gordon |publisher=Ten Speed Press |year= 1998 |isbn=978-0-89815-977-6}}</ref> The [[Torah]] states the only kosher flying insects with four walking legs have knees that extend above their feet so that they hop.<ref>{{citation |url= http://bible.ort.org/books/pentd2.asp?action=displayanchor&pentid=P3008 |title=Navigating the Bible: Leviticus }}</ref><!-- Thus nonjumping Orthoptera such as [[mole crickets]] are certainly not kosher.-->

===As creators of biofuel===

With new research showing promise in locating alternative [[biofuel]] sources in the gut of insects, grasshoppers are one species of interest. The insect's ability to break down [[cellulose]] and [[lignin]] without producing [[greenhouse gases]] has aroused scientific interest.<ref>{{Cite journal |last1=Shi |first1=Weibing |last2=Xie |first2=Shangxian |last3=Chen |first3=Xueyan |last4=Sun |first4=Su |last5=Zhou |first5=Xin |last6=Liu |first6=Lantao |last7=Gao |first7=Peng |last8=Kyrpides |first8=Nikos C. |last9=No |first9=En-Gyu |date=January 2013 |title=Comparative Genomic Analysis of the Endosymbionts of Herbivorous Insects Reveals Eco-Environmental Adaptations: Biotechnology Applications |journal=PLOS Genetics |volume=9 |issue=1 |pages=e1003131 |doi=10.1371/journal.pgen.1003131 |pmc=3542064 |pmid=23326236 |doi-access=free }}</ref>

==See also==
* [[List of Orthopteroid genera containing species recorded in Europe]]
* [[List of Orthoptera recorded in Britain]]
* [[Orthopterida]]
* [[Female sperm storage]]

==References==
{{Reflist}}

== External links ==
{{Commons category}}
{{Wikispecies}}
*[https://orthoptera.speciesfile.org/ Orthoptera Species File Online]
*[https://www.ent.iastate.edu/imagegal/orthoptera/ Orthoptera Image Gallery (Iowa State University Entomology Department)]
*[https://web.archive.org/web/20050615144819/http://www.affa.gov.au/content/output.cfm?ObjectID=D2C48F86-BA1A-11A1-A2200060B0A00383 Australian Plague Locust Commission]
*[https://orthsoc.org/ The Orthopterists' Society]
*[http://www.acrida.info AcridAfrica, les acridiens d'Afrique de l'Ouest]
*{{Cite NIE|wstitle=Orthoptera|year=1905 |short=x}}
*[https://web.archive.org/web/20130722161334/http://ricochetscience.com/birdwing_grasshopper/ Birdwing Grasshoppers in Belize]
*[https://bio.acousti.ca/classification/orthoptera Sound recordings of Orthoptera at BioAcoustica]

{{Orders of Insects}}
{{Orthoptera}}

{{Taxonbar|from=Q167810}}
{{Authority control}}

[[Category:Orthoptera| ]]
[[Category:Insect orders]]
[[Category:Edible insects]]
[[Category:Carboniferous first appearances]]
[[Category:Taxa named by Pierre André Latreille]]
[[Category:Orthopterida]]