{{Short description|Clade of wingless insects}} {{For|the plant genus|Notoptera (plant)}} {{Automatic taxobox | fossil_range = {{fossilrange|320|0}} Late Carboniferous-Present | image = Mantophasma zephyra Zompro et al 2002.jpg | image_caption = ''Mantophasma zephyra'' (Mantophasmatidae) | image2 = Grylloblattidae (cropped).jpg | image2_caption = Member of Grylloblattidae | taxon = Notoptera | authority = Crampton, 1915 | subdivision_ranks = Subgroups & families | subdivision = *{{extinct}}Camptoneuritidae *{{extinct}}Tillyardembiidae *{{extinct}}Geinitziidae *'''Grylloblattodea''' **{{extinct}}''Aristovia'' **{{extinct}}Blattogryllidae **{{extinct}}Plesioblattogryllidae **{{extinct}}''Zygogrylloblatta'' **Grylloblattidae (ice-crawlers) *'''Mantophasmatodea''' **Mantophasmatidae (gladiators) And see text | synonyms = Xenonomia <small>Terry & Whiting, 2005</small> }}
'''Notoptera''', also known as '''Xenonomia''',<ref name=":0">{{Cite journal |last=Cui |first=Yingying |last2=Bardin |first2=Jérémie |last3=Wipfler |first3=Benjamin |last4=Demers‐Potvin |first4=Alexandre |last5=Bai |first5=Ming |last6=Tong |first6=Yi‐Jie |last7=Chen |first7=Grace Nuoxi |last8=Chen |first8=Huarong |last9=Zhao |first9=Zhen‐Ya |last10=Ren |first10=Dong |last11=Béthoux |first11=Olivier |date=2024-03-07 |title=A winged relative of ice‐crawlers in amber bridges the cryptic extant Xenonomia and a rich fossil record |url=https://onlinelibrary.wiley.com/doi/10.1111/1744-7917.13338 |journal=Insect Science |language=en |doi=10.1111/1744-7917.13338 |issn=1672-9609|url-access=subscription }}</ref> is a clade of insects belonging to Polyneoptera. It contains two living groups, Mantophasmatidae (gladiators) native to southern Africa, and Grylloblattidae (ice crawlers) native to cold montane environments in the Northern Hemisphere. Both groups are wingless.
==History of research== The name was originally coined in 1915 as an order for Grylloblattidae and largely forgotten until it was resurrected and redefined ("Notoptera Crampton ''sensu novum''") by Engel and Grimaldi in 2004 (after the discovery of living Mantophasmatidae), who recommended to give a single order that includes both the living and fossil representatives of the lineage.<ref name="Engel Grimaldi 2004">{{cite journal | last1=Engel | first1=Michael S. | last2=Grimaldi| first2=David A. | title=A New Rock Crawler in Baltic Amber, with Comments on the Order(Mantophasmatodea: Mantophasmatidae) | journal=American Museum Novitates | publisher=American Museum of Natural History (BioOne sponsored) | issue=3431 | year=2004 | pages=1–12 | issn=0003-0082 | doi=10.1206/0003-0082(2004)431<0001:anrcib>2.0.co;2 | s2cid=85653533 | url=https://zenodo.org/record/5375006 }}</ref>
Terry and Whiting in 2005 independently proposed a new name, "Xenonomia", for the same lineage of insects (including the Grylloblattodea and Mantophasmatodea, treated as orders).<ref name="Terry Whiting 2005 pp. 240–257">{{cite journal | last1=Terry | first1=Matthew D. | last2=Whiting | first2=Michael F. | title=Mantophasmatodea and phylogeny of the lower neopterous insects | journal=Cladistics | publisher=Wiley | volume=21 | issue=3 | year=2005 | issn=0748-3007 | doi=10.1111/j.1096-0031.2005.00062.x | pages=240–257| s2cid=86259809 | doi-access=free }}</ref> The orders Grylloblattodea and Mantophasmatodea are sometimes ranked as suborders of a single order, Notoptera.<ref name=ArilloEngel2006>{{cite journal|last1=Arillo|first1=A.|last2=Engel|first2=M.S.|year=2006|title=Rock crawlers in Baltic amber (Notoptera: Mantophasmatodea)|journal=American Museum Novitates|issue=3539|pages=1–10|doi=10.1206/0003-0082(2006)3539[1:RCIBAN]2.0.CO;2 |url=https://www.biodiversitylibrary.org/page/59821931}}</ref> Some authors consider this the valid name of the group.<ref name=":0" />
== Evolutionary history == [[File:Peng et al. 2025 f04.jpg|thumb|Life restoration of ''Zygogrylloblatta'', a winged "grylloblattidan" from the mid-Cretaceous|left]] The earliest stem-group representatives of Notoptera/Xenonomia had emerged by the Late Carboniferous, around 320 million years ago. Early members of the group, which unlike modern notopterans had wings, have been referred to as members of "Grylloblattida" and "Reculida", with their relationships to modern notopterans historically being the subject of controversy. Winged "grylloblattidans" reached their apex of diversity during the Permian (299-252 million years ago), where they represented up to a third of all insects at some localities.<ref name=":0" /> The earliest mantophasmatids are known from the Middle Jurassic of China, around 160 million years ago.<ref>{{Cite journal |last1=Huang |first1=Di-ying |last2=Nel |first2=André |last3=Zompro |first3=Oliver |last4=Waller |first4=Alain |date=2008-06-11 |title=Mantophasmatodea now in the Jurassic |url=https://www.researchgate.net/publication/5311278 |journal=Naturwissenschaften |volume=95 |issue=10 |pages=947–952 |doi=10.1007/s00114-008-0412-x |issn=0028-1042 |pmid=18545982 |s2cid=35408984}}</ref> No fossil record of modern grylloblattids is known, though the winged ''Aristovia'' and ''Zygogrylloblatta'' are known from the Burmese amber of Myanmar dating to the mid-Cretaceous around 100 million years ago, have mouthparts very similar to modern grylloblattids, indicating their closer relationship to modern grylloblattids than to mantophasmatids.<ref name=":0" /><ref name=":1">{{Cite journal |last=Peng |first=Ancheng |last2=Engel |first2=Michael S. |last3=Boderau |first3=Mathieu |last4=Legendre |first4=Frédéric |last5=Liu |first5=Yu |last6=Nyunt |first6=Thet Tin |last7=Wang |first7=Bo |last8=Nel |first8=André |date=June 2025 |title=Descending from trees: a Cretaceous winged ice-crawler illuminates the ecological shift and origin of Grylloblattidae |url=https://royalsocietypublishing.org/doi/10.1098/rspb.2025.0557 |journal=Proceedings of the Royal Society B: Biological Sciences |language=en |volume=292 |issue=2049 |doi=10.1098/rspb.2025.0557 |issn=1471-2954|pmc=12173512 }}</ref> Mantophasmatids and grylloblattids are thought to have lost their wings independently. Winged "grylloblattidans" declined in diversity and abundance from the Triassic onwards, with the youngest records of winged "grylloblattidans" dating to the Early-mid Cretaceous.<ref name=":0" />
== List of extinct families == Following Aristov (2015):<ref>{{cite journal|last=Aristov|first=D. S.|year=2015|title=Classification of the order Eoblattida (Insecta: Blattidea) with description of new taxa|journal=Far Eastern Entomologist|issue=301|pages=1–56|url=https://www.biosoil.ru/storage/entities/fscpublication/478/85f3310d-8744-4f28-ad6a-95da694b877f.pdf|access-date=1 November 2025}}</ref>
'''Eoblattida'''
* †Atactophlebiidae * †Bardapteridae * †Blattogryllidae * †Cacurgidae? (other authors place this family in Panorthoptera and unrelated to Notoptera<ref name="Schnubeletal2019">{{cite journal |author1=Thomas Schubnel |author2=Dawn Roberts |author3=Patrick Roques |author4=Romain Garrouste |author5=Laure Desutter-Grandcolas |author6=André Nel |year=2019 |title=Moscovian fossils shed light on the enigmatic polyneopteran families Cacurgidae and Eoblattidae (Insecta: 'Eoblattida', Archaeorthoptera) |journal=Journal of Systematic Palaeontology |volume=18 |issue=6 |pages=499–511 |doi=10.1080/14772019.2019.1627595 |s2cid=203899153 |url=https://hal.sorbonne-universite.fr/hal-02294662/file/Schubnel%20et%20al.%20-%202019%20-%20Moscovian%20fossils%20shed%20light%20on%20the%20enigmatic%20poly.pdf }}</ref>) * †Daldubidae * †Doubraviidae * †Eoblattidae? (other authors place this family in Archaeorthoptera and unrelated to Notoptera<ref name=":1" />) * †Euryptilonidae * †Ideliidae? (other authors place this family in Paoliida as a synonym of Paoliidae and unrelated to Notoptera<ref name="Prokopetal2014">{{Cite journal |author1=Jakub Prokop |author2=Wieslaw Krzemiński |author3=Ewa Krzemińska |author4=Thomas Hörnschemeyer |author5=Jan-Michael Ilger |author6=Carsten Brauckmann |author7=Philippe Grandcolas |author8=André Nel |year=2014 |title=Late Palaeozoic Paoliida is the sister group of Dictyoptera (Insecta: Neoptera) |journal=Journal of Systematic Palaeontology |volume=12 |issue=5 |pages=601–622 |doi=10.1080/14772019.2013.823468|s2cid=84407734 }}</ref><ref name="NelPoschmann2021">{{cite journal|author1=André Nel|author2=Markus J. Poschmann |year=2021|title=Comparison of the recently described early Permian paoliid genus ''Permomertovia'' with the 'eoblattid' genera of the families Permulidae and Permotermopsidae|journal=Palaeoentomology|volume=4|issue=1|pages=39–43|doi=10.11646/palaeoentomology.4.1.8|url=https://tb.plazi.org/GgServer/summary/6C58FF98FFBC291D117D2215FFF3FFA3|url-access=subscription}}</ref>) * †Idelinellidae * †Megakhosaridae * †Mesorthopteridae * †Permopectinidae * †Permotermopsidae? (other authors place this family in Paoliida as a synonym of Paoliidae and unrelated to Notoptera<ref name="NelPoschmann2021"/>) * †Protophasmatidae * †Soyanopteridae * †Tococladidae? (other authors place this family in Cnemidolestodea and unrelated to Notoptera<ref>{{Cite journal|author1=Jakub Prokop |author2=Jacek Szwedo |author3=Jean Lapeyrie |author4=Romain Garrouste |author5=André Nel |year=2015 |title=New Middle Permian insects from Salagou Formation of the Lodève Basin in southern France (Insecta: Pterygota) |journal=Annales de la Société Entomologique de France |series=New Series |volume=51 |issue=1 |pages=14–51 |doi=10.1080/00379271.2015.1054645 |s2cid=87501340 }}</ref>)
'''Reculida'''
* †Archiprobnidae * †Bajanzhargalanidae * †Camptoneuritidae * †Chaulioditidae * †Chelopteridae * †Epideigmatidae * †Euremiscidae * †Geinitziidae * †Gorochoviidae * †Havlatiidae * †Ivapteridae * †Kargalopteridae * †Lemmatophoridae * †Liomopteridae * †Madygenophlebiidae * †Neleidae * †Probnidae * †Protoblattinidae? (other authors place this family in Paoliida as a synonym of Paoliidae and unrelated to Notoptera<ref name="Prokopetal2014"/>) * †Reculidae * †Sinonamuropteridae * †Skaliciidae * †Sylvaphlebiidae
==Gallery== <gallery widths="220"> File:GrylloblattaCampodeiformis.jpg|''Grylloblatta campodeiformis'' File:Gladiator.jpg|A nymph of a mantophasmatid </gallery>
==See also== * Grylloblattidae * Mantophasmatidae * Palaeocixiidae
==Further reading== *Ando H. 1982. ''Biology of the Notoptera''. Kashiyo-Insatsu Co. Ltd., Nagano, Japan.
==References== {{Reflist}}
==External links== *{{commons category-inline|Notoptera}} *{{Wikispecies-inline|Notoptera}}
{{Orders of Insects}} {{Notoptera}} {{Taxonbar|from=Q1185781}}
Category:Notoptera Category:Extant Triassic first appearances Category:Polyneoptera