View source for Ensifera

{{Short description|Suborder of cricket-like animals}}
{{For|the avian genus ''Ensifera''|Sword-billed hummingbird}}
{{Automatic taxobox
|fossil_range = [[Artinskian]]/[[Kungurian]]&ndash;[[Holocene]], {{fossilrange|272.3|0|earliest=290.1|ref=<ref name="Wang2016">{{cite journal | last1 = Wang | first1 = Yan-hui | last2 = Engel | first2 = Michael S. | last3 = Rafael | first3 = José A. | last4 = Wu | first4 = Hao-yang | last5 = Rédei | first5 = Dávid | last6 = Xie | first6 = Qiang | last7 = Wang | first7 = Gang | last8 = Liu | first8 = Xiao-guang | last9 = Bu | first9 = Wen-jun | date = 2016 | title = Fossil record of stem groups employed in evaluating the chronogram of insects (Arthropoda: Hexapoda) | journal = [[Scientific Reports]] | volume = 6 | page = 38939 | doi = 10.1038/srep38939 | pmid=27958352 | pmc=5154178| bibcode = 2016NatSR...638939W }}</ref>}}
| image = Katydid.jpg
| image_caption = A [[Tettigoniidae|bush cricket or katydid]]
| taxon = Ensifera
| authority =
| subdivision_ranks = Superfamilies and families
| subdivision = See text
}}

'''Ensifera''' is a [[Order (biology)|suborder]] of [[insect]]s that includes the various types of crickets and their allies, including [[cricket (insect)|true crickets]], [[Rhaphidophoridae|camel crickets]], [[Tettigoniidae|bush crickets or katydid]]s, [[Prophalangopsidae|grig]]s, ''[[wētā]]'', and [[Cooloola (insect)|Cooloola monster]]s. 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,<ref name=Resh>{{cite book |author1=Resh, Vincent H.|author2=Cardé, Ring T. |title=Encyclopedia of Insects |url=https://books.google.com/books?id=Jk0Hym1yF0cC&q=Grylloidea |year=2009 |publisher=Academic Press |isbn=978-0-08-092090-0 |pages=232, 733–735}}</ref> the split having occurred at the end of the [[Permian]] period.<ref>Zeuner, F. E. (1939). ''Fossil Orthoptera Ensifera''. London: British Museum Natural History.</ref> Unlike the Caelifera, the Ensifera contain numerous members that are partially carnivorous, feeding on other insects, as well as plants.

''Ensifer'' is [[Latin]] for "sword bearer", and refers to the typically elongated and blade-like [[ovipositor]] of the females.<ref>{{cite web |url=http://www.nhm.ac.uk/research-curation/life-sciences/terrestrial-invertebrates/organisms/insects/orthopteroid-insects/orthopteriod-orders/orthoptera/index.html |title=Orthoptera |publisher=Natural History Museum |access-date=2015-05-13}}</ref>

==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.<ref name=Resh/>

Ensiferans are distinguished from Caeliferans by their elongated, threadlike [[Antenna (biology)|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 organ]]s are located on the tibiae of the front legs.<ref name=TOL/> 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 [[protein]]aceous [[spermatophylax]], the function of which is to provide a nutritional [[nuptial gift]] to the female.<ref name=TOL>{{cite web |url=http://tolweb.org/Ensifera/13315 |title=Ensifera: Crickets, katydids and weta |author1=Gwynne, Darryl T. |author2=DeSutter |author3=Laure |year=1996 |publisher=TOLweb |access-date=8 May 2015}}</ref><ref>{{cite journal | author1 = Vahed | author2 = K. | year = 1998 | title = The function of nuptial feeding in insects: review of empirical studies | journal = Biological Reviews | volume = 73 | pages = 43–78 | doi = 10.1111/j.1469-185X.1997.tb00025.x | s2cid = 86644963 | url = http://www.famu.org/mayfly/pubs/zor/zorpub_vahedk1998p45.pdf | access-date = 2015-05-08 | archive-url = https://web.archive.org/web/20120311042845/http://www.famu.org/mayfly/pubs/zor/zorpub_vahedk1998p45.pdf | archive-date = 2012-03-11 | url-status = dead }}</ref>

==Taxonomy==
[[File:Mole cricket02.jpg|thumb|upright|A [[mole cricket]], showing the front legs specialised for digging]]
[[File:Ceuthophiluscricket.jpg|thumb|upright|A [[Rhaphidophoridae|cave cricket]], showing the long hind legs and antennae]]
[[File:CSIRO ScienceImage 10713 Cooloola Monster.jpg|thumb|upright|A [[Cooloola monster]], a subterranean family from [[Queensland]], Australia]]
[[File:Пальцепалый кузнечик.jpg|thumb|upright|A [[Schizodactylidae|splay-footed cricket]] from South India]]

The Orthoptera Species File database lists the following superfamilies and families.<ref>{{cite web |url= http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1121518 |title=Suborder Ensifera |work=Orthoptera Species File |access-date=9 May 2015}}</ref>
*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]]'' {{small|Heer, 1865}}
*Infraorder [[Gryllidea]]
**Superfamily [[Grylloidea]] 
***Family †[[Baissogryllidae]] 
***Family [[Gryllidae]] - true crickets
***Family [[Mogoplistidae]] - scaly crickets
***Family [[Phalangopsidae]] {{small|Blanchard, 1845}}
***Family †[[Protogryllidae]] {{small|Zeuner, 1937}}
***Family [[Trigonidiidae]] {{small|Saussure, 1874}}
**Superfamily [[Gryllotalpoidea]] {{small|Leach, 1815}}<ref>[http://orthoptera.speciesfile.org/Common/basic/Taxa.aspx?TaxonNameID=1128272 Orthoptera Species File (Version 5.0/5.0)]</ref>
***Family [[Gryllotalpidae]] {{small|Leach, 1815}} – mole crickets
***Family [[Myrmecophilidae]] {{small|Saussure, 1874}} - ant crickets
*Infraorder †[[Oedischiidea]] 
**Superfamily †[[Oedischioidea]] {{small|Handlirsch, 1906}}
***Family †[[Anelcanidae]] {{small|Carpenter, 1986}}
***Family †[[Bintoniellidae]] {{small|Handlirsch, 1939}}
***Family †[[Mesoedischiidae]] {{small|Gorochov, 1987}}
***Family †[[Oedischiidae]] {{small|Handlirsch, 1906}}
***Family †[[Proparagryllacrididae]] {{small|Riek, 1956}}
***Family †[[Pruvostitidae]] {{small|Zalessky, 1929}}
***Family ''incertae sedis''
****Genus †''[[Crinoedischia]]'' {{small|Béthoux & Beckemeyer, 2007}}
****Genus †''[[Loxoedischia]]'' {{small|Beckemeyer, 2011}}
**Superfamily †[[Triassomantoidea]] {{small|Tillyard, 1922}}
***Family †[[Adumbratomorphidae]] {{small|Gorochov, 1987}}
***Family †[[Triassomantidae]] {{small|Tillyard, 1922}}
**Superfamily †[[Xenopteroidea]] {{small|Riek, 1955}}
***Family †[[Xenopteridae]] {{small|Riek, 1955}}
**Superfamily ''incertae sedis''
***family †[[Permotettigoniidae]] {{small|Nel & Garrouste, 2016}}
***Family ''incertae sedis''
****Genus †''[[Permophyllum]]'' {{small|Prokop, ''et al'', 2015}}
*Infraorder [[Tettigoniidea]] 
**Superfamily [[Hagloidea]] - grigs
***Family †[[Eospilopteronidae]] {{small|Cockerell, 1916}}
***Family †[[Haglidae]] {{small|Handlirsch, 1906}}
***Family †[[Hagloedischiidae]] {{small|Gorochov, 1986}}
***Family †[[Prezottophlebiidae]] {{small|Martins-Neto, 2007}}
***Family [[Prophalangopsidae]] {{small|Kirby, 1906}}
***Family †[[Tuphellidae]] {{small|Gorochov, 1988}}
**Superfamily †[[Phasmomimoidea]] {{small|Sharov, 1968}}
***Family †[[Phasmomimidae]] {{small|Sharov, 1968}}
**Superfamily [[Rhaphidophoroidea]] {{small|Walker, 1869}}
***Family [[Rhaphidophoridae]] {{small|Walker, 1869}} - camel crickets, cave crickets, cave weta
**Superfamily [[Schizodactyloidea]] {{small|Blanchard, 1845}}
***Family [[Schizodactylidae]] {{small|Blanchard, 1845}} - dune or splay-footed crickets
**Superfamily [[Stenopelmatoidea]] {{small|Burmeister, 1838}}
***Family [[Anostostomatidae]] {{small|Saussure, 1859}} - weta (except cave weta), king crickets
***Family [[Cooloolidae]] {{small|Rentz, 1980}} - Cooloola monsters
***Family [[Gryllacrididae]] {{small|Blanchard, 1845}} - leaf-rolling crickets
***Family  [[Stenopelmatidae]] {{small|Burmeister, 1838}} - Jerusalem crickets
**Superfamily [[Tettigonioidea]] {{small|Krauss, 1902}}
***Family †[[Haglotettigoniidae]] {{small|Gorochov, 1988}}
***Family [[Tettigoniidae]] <small>Krauss, 1902</small> - bush crickets, katydids, koringkrieks
**Superfamily ''Incertae sedis''
***Family ''incertae sedis''
****Genus †''[[Tettoraptor]]'' {{small|Gorochov, 2012}}
*Infraorder ''incertae sedis''
**Superfamily †[[Gryllavoidea]] {{small|Gorochov, 1986}} 
***Family †[[Gryllavidae]] {{small|Gorochov, 1986}}
*Superfamily ''Incertae sedis''
** Family †[[Palaeorehniidae]] (syn "[[Zeuneropterinae]]")<ref name=Republicopteron>{{Cite journal |last1=Archibald |first1=S. B. |last2=Gu |first2=J.-J. |last3=Mathewes |first3=R. W. |title=The Palaeorehniidae (Orthoptera, Ensifera, "Zeuneropterinae"), and new taxa from the early Eocene Okanagan Highlands, western North America |year=2022 |journal=Zootaxa |volume=5100 |issue=4 |pages=559–572 |doi=10.11646/zootaxa.5100.4.6 |pmid=35391059 }}</ref>
**Family †[[Vitimiidae]]

==Phylogeny==
[[File:Pycnophlebia speciosa.JPG|thumb|[[Prophalangopsidae]]: [[Jurassic]] fossil ''[[Pycnophlebia speciosa]]'']]

The [[phylogeny|phylogenetic relationships]] of the Ensifera, summarized by Darryl Gwynne in 1995 from his own work and that of earlier authors,{{efn|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.<ref name=Gwynne/>}} 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.<ref name=TOL/><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>

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.<ref>{{cite journal |last1=Prokop |first1=Jakub |last2=Nel |first2=André |last3=Hoch |first3=Ivan |title=Discovery of the oldest known Pterygota in the Lower Carboniferous of the Upper Silesian Basin in the Czech Republic (Insecta: Archaeorthoptera) |journal=Geobios |date=2005 |volume=38 |issue=3 |pages=383–387 |doi=10.1016/j.geobios.2003.11.006|bibcode=2005Geobi..38..383P }}</ref>

{{clade| style=font-size:85%;line-height:85%
 |label1=[[Orthoptera]]
 |1={{clade
    |label1='''Ensifera'''
    |1={{clade
        |label1=Elcanidea
        |1={{clade
            |1=†[[Elcanoidea]]
            |2=†[[Permoraphidioidea]]
            }}
        |label2=Oedischiidea
        |2={{clade
            |1=†[[Oedischioidea]]
            |2=†[[Triassomantoidea]]
            |3=†[[Xenopteroidea]]
            }}
        |label3=Tettigoniidea
        |3={{clade
            |1=[[Hagloidea]]: (including grigs)
            |2=†[[Phasmomimoidea]]
            |3=[[Stenopelmatoidea]] (weta, king crickets)
            |4=[[Tettigonioidea]] (bush crickets, katydids, koringkreiks)
            }}
        |4=[[Rhaphidophoroidea]] (cave weta, cave crickets)
        |5=†[[Gryllavoidea]]
        |6=[[Grylloidea]] (crickets)
        |7=[[Schizodactyloidea]] (dune crickets)
        }}
     |label2=[[Caelifera]]
     |2=([[Acrididea|grasshoppers]], [[Tetrigidae|groundhoppers]], [[Tridactyloidea|pygmy mole crickets]])
    }}
}}

==Notes==
{{notelist}}

==References==
{{Reflist|28em}}

==External links==
* [http://www.orthoptera.org/ The Orthopterists' Society]

{{Orthoptera|1}}
{{Taxonbar|from=Q27736}}
{{Authority control}}

[[Category:Ensifera| ]]
[[Category:Insect suborders]]
[[Category:Cisuralian first appearances]]
[[Category:Extant Permian first appearances]]