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{{Short description|Suborder of cricket-like animals}}
{{for|the avian genus Ensifera|Sword-billed Hummingbird}}
{{For|the avian genus ''Ensifera''|Sword-billed hummingbird}}
{{Automatic taxobox
{{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>}}
|fossil_range= [[Carboniferous]] - [[Recent]] {{fossilrange|395|0}}
| image = Katydid.jpg
| image = Katydid.jpg
| image_caption = Katydid
| image_caption = A [[Tettigoniidae|bush cricket or katydid]]
| subordo = Ensifera
| taxon = Ensifera
| authority =
| subdivision_ranks = [[Taxonomic rank|Superfamilies]] and [[family (biology)|families]]
| subdivision_ranks = Superfamilies and families
| subdivision =
see text
| 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|weta]] 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.
'''Ensifera''' is a suborder of the order [[Orthoptera]], comprising [[insect]]s commonly known as [[Cricket (insect)|cricket]]s (or [[Tettigoniidae|bush crickets]]) and [[Tettigoniidae|katydids]]. "Ensifer" means "sword bearer" in [[Latin]], and refers to the typically elongated and blade-like [[ovipositor]] of the females. The classification of the [[suborder]] was drastically revised in December 2005 by the [[Orthopterists' Society]].


''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>
==Classification==

*Superfamily [[Grylloidea]]
==Characteristics==
**[[Gryllidae]] - true crickets
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/>
**[[Gryllotalpidae]] - mole crickets

**[[Mogoplistidae]]
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>
**[[Myrmecophilidae]] - ant crickets

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

*Superfamily [[Stenopelmatoidea]]
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>
**[[Anostostomatidae]] - wetas, king crickets
*Infraorder †[[Elcanidea]]
**[[Cooloolidae]]
**Superfamily †[[Elcanoidea]]
**[[Gryllacrididae]] - leaf-rolling crickets
***Family †[[Elcanidae]] (Late Triassic - Paleocene)
**[[Stenopelmatidae]] - Jerusalem crickets
***Family †[[Permelcanidae]] (Early Permian - Late Triassic)
*Superfamily [[Tettigonioidea]] (or [[Tettigoniidae]]) - katydids, koringkrieks
**Superfamily †[[Permoraphidioidea]]
**[[Haglotettigoniidae]]
***Family †[[Permoraphidiidae]] (Permian)
**[[Tettigoniidae]]
***Family †[[Pseudelcanidae]] (Early Permian)
*Ensifera incertae sedis
***Family †[[Thueringoedischiidae]] (Early Permian)
**[[Elcanidea]]
***Family ''[[incertae sedis]]''
**[[Oedischiidea]]
****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==
==External links==
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{{Orthoptera|1}}
{{Orthoptera|1}}
{{Taxonbar|from=Q27736}}
{{Authority control}}


[[Category:Orthoptera]]
[[Category:Ensifera| ]]
[[Category:Insect suborders]]

[[Category:Cisuralian first appearances]]
{{Orthoptera-stub}}
[[Category:Extant Permian first appearances]]

[[ja:バッタ目#キリギリス亜目（剣弁亜目） Ensifera]]

Latest revision as of 14:51, 15 August 2024

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

Ensifera
Temporal range: Artinskian/Kungurian–Holocene, 272.3–0 Ma[1]
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

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.

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

Characteristics

[edit]

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

[edit]
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]

Phylogeny

[edit]
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,[a] 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
Oedischiidea
Tettigoniidea

Hagloidea: (including grigs)

†Phasmomimoidea

Stenopelmatoidea (weta, king crickets)

Tettigonioidea (bush crickets, katydids, koringkreiks)

Rhaphidophoroidea (cave weta, cave crickets)

†Gryllavoidea

Grylloidea (crickets)

Schizodactyloidea (dune crickets)

Caelifera

(grasshoppers, groundhoppers, pygmy mole crickets)

Notes

[edit]
  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

[edit]
  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. Bibcode:2016NatSR...638939W. doi:10.1038/srep38939. PMC 5154178. PMID 27958352.
  2. ^ a b 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. ^ a b c 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. PMID 35391059.
  10. ^ a b 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. Bibcode:2005Geobi..38..383P. doi:10.1016/j.geobios.2003.11.006.
[edit]
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