The wingless insect order Notoptera, a group first proposed in 1915, has been largely unused since its original conception, but in the most recent classification of the lineage of insects that includes the Grylloblattodea and Mantophasmatodea,[1] the name was resurrected and redefined so as to give a single order that includes both the living and fossil representatives of the lineage.

Temporal range: 252–0 Ma
Triassic - Recent
Mantophasma zephyra Zompro et al 2002
Mantophasma zephyra
Scientific classification

Crampton, 1915
Suborders & families
  • Family Tillyardembiidae
  • Family † Blattogryllidae

Suborder Grylloblattodea

Suborder Mantophasmatodea


As now defined, the order comprises four families, two of them known only from fossils, two known from both fossil and living representatives, and fewer than 60 known species in total.



A nymph of a mantophasmatid

See also


  1. ^ Arillo, A. & M. Engel (2006) Rock Crawlers in Baltic Amber (Notoptera: Mantophasmatodea). American Museum Novitates 3539:1-10

External links


Bajanzhargalanidae is an extinct family of rock crawlers in the order Grylloblattodea. There are at least four genera and about seven described species in Bajanzhargalanidae.


The Dicondylia are a taxonomic group (taxon) that includes all insects except the jumping bristletails (Archaeognatha). Dicondylia have a mandible attached with two hinges to the head capsule (dicondyl), in contrast to the original mandible with a single ball joint (monocondyl).


Eumetabola is an unranked category of Neoptera. Two large unities known as the Paurometabola and Eumetabola are probably from the adelphotaxa of the Neoptera after exclusion of the Plecoptera. The monophyly of these unities appears to be weakly justified.


The Exopterygota, also known as Hemipterodea, are a superorder of insects of the subclass Pterygota in the infraclass Neoptera, in which the young resemble adults but have externally developing wings. They undergo a modest change between immature and adult, without going through a pupal stage. The nymphs develop gradually into adults through a process of moulting.

The Exopterygota are a highly diverse insect superorder, with at least 130,000 living species divided between 15 orders. They include cockroaches, termites, grasshoppers, thrips, lice and stick insects, among many other types of insects.

They are distinguished from the Endopterygota (or Holometabola) by the way in which their wings develop. Endopterygota (meaning literally "internal winged forms") develop wings inside the body and undergo an elaborate metamorphosis involving a pupal stage. Exopterygota ("external winged forms") develop wings on the outside of their bodies without going through a true pupal stage, though a few have something resembling a pupa (e.g., Aleyrodidae).

Ephemeroptera (mayflies) and Odonata (dragonflies and damselflies) also have gradual wing development, this being a plesiomorphic trait. These two orders belong to the infraclass Palaeoptera however, which is not included in Neoptera. As opposed to Neoptera, they cannot fold their wings over their back in the horizontal plane, only vertically (as damselflies do) if at all.


Gorochoviidae is an extinct family of rock crawlers in the order Grylloblattodea. There are at least three genera and about eight described species in Gorochoviidae.

Grylloblatta bifratrilecta

Grylloblatta bifratrilecta is a species of rock crawler in the family Grylloblattidae. It is found in North America, including Sonora Pass in the Sierra Nevada. It lives in crevices at high altitudes, typically above 7,000 feet. It is nocturnal and active mostly during winter months.

Grylloblatta chintimini

Grylloblatta chintimini is a species of rock crawler in the family Grylloblattidae. It is found in North America.


Grylloblattidae, the icebugs, or ice crawlers, is a family of extremophile (psychrophile) and wingless insects that live in the cold on top of mountains and the edges of glaciers. They belong, along with Mantophasmatidae (rock crawlers), to the order Notoptera. Grylloblattids are tiny, wingless insects with a head resembling that of a cockroach, with long antennae and having elongated cerci arising from the tip of their abdomen. They cannot tolerate warmth (most species will die at 10° C) and many species have small distribution ranges.


Madygenophlebiidae is an extinct family of rock crawlers in the order Grylloblattodea. There are at least two genera and four described species in Madygenophlebiidae.


Mantophasmatidae is a family of carnivorous wingless insects within the order Notoptera, which was discovered in Africa in 2001. Originally, the group was regarded as an order in its own right, and named Mantophasmatodea, but based on recent evidence indicating a sister group relationship with Grylloblattidae (formerly classified in the order Grylloblattodea), Arillo & Engel have combined the two groups into a single order, Notoptera.


Neoptera is a classification group that includes most orders of the winged insects, specifically those that can flex their wings over their abdomens. This is in contrast with the more basal orders of winged insects (the "Palaeoptera" assemblage), which are unable to flex their wings in this way.


The Orthopterida is a superorder of the Polyneoptera that represents the extant orders Orthoptera (grasshoppers, crickets, and katydids), and Phasmatodea (stick insects and leaf insects). The Orthopterida also includes the extinct orders Titanoptera and Caloneurodea. There is general consensus of monophyly in this superorder, based on reduction of the second valvulae, an ovipositor derived from the gonoplac, and an enlarged precostal region on the forewing.

The two other superorders of the Polyneoptera are the Plecopterida, which represents the orders Plecoptera (stoneflies), Emboidea (Embioptera/Embiidina; webspinners), and Zoraptera (angel insects), and the Dictyoptera, which represents Blattodea (cockroaches & termites), and Mantodea (mantids). Two other orders, the Notoptera (ice-crawlers and gladiators) and Dermaptera (earwigs) are also placed in the Polyneoptera but outside the superorders discussed above.


Otopappus is a genus of flowering plants in the sunflower tribe within the daisy family, primarily Mesoamerican but with one species from Jamaica.

SpeciesOtopappus acuminatus S.Watson - Jalisco

Otopappus brevipes B.L.Rob. - from Chiapas to Nicaragua

Otopappus calarcanus S.Díaz - Colombia

Otopappus curviflorus (R.Br.) Hemsl. - from Veracruz to Nicaragua

Otopappus epaleaceus Hemsl. - from Puebla to Guatemala

Otopappus glabratus (J.M.Coult.) S.F.Blake - Guatemala, Honduras, El Salvador

Otopappus guatemalensis (Urb.) R.L.Hartm. & Stuessy - Guatemala, Belize, Yucatán

Otopappus hirsutus (Sw.) R.L.Hartm. & Stuessy - Jamaica

Otopappus imbricatus (Sch.Bip.) S.F.Blake - Puebla, Morelos, Michoacán, Guerrero

Otopappus koelzii McVaugh - Michoacán, Colima, Jalisco

Otopappus mexicanus (Rzed.) H.Rob. - Guerrero, Oaxaca

Otopappus microcephalus S.F.Blake - Colima, Jalisco, Guerrero, Oaxaca

Otopappus robustus Hemsl. - Veracruz

Otopappus scaber S.F.Blake - Guatemala, Belize, Yucatán, Chiapas

Otopappus syncephalus Donn.Sm. - Guatemala

Otopappus tequilanus (A.Gray) B.L.Rob. - Sinaloa, Jalisco, Nayarit, Michoacán, Guerrero, Oaxaca

Otopappus verbesinoides Benth. - from Oaxaca to Nicaraguaformerly includedSeveral species one included in Otopappus are now regarded as better suited to other genera: Lundellianthus Oblivia Verbesina


Palaeocixiidae is an extinct family of rock crawlers in the order Grylloblattodea. There is one genus, Palaeocixius, in Palaeocixiidae.


Panorpida or Mecopterida is a proposed superorder of Endopterygota. The conjectured monophyly of the Panorpida is historically based on morphological evidence, namely the reduction or loss of the ovipositor and several internal characteristics, including a muscle connecting a pleuron and the first axillary sclerite at the base of the wing, various features of the larval maxilla and labium, and basal fusion of CuP and A1 veins in the hind wings. The monophyly of the Panorpida is also supported by recent molecular data.


The cohort Polyneoptera is probably the most appropriate taxonomic ranking for commonly-used terms such as Orthopteroid insects: namely the orders similar to the Orthoptera (grasshoppers, crickets, etc.). They are all winged insects (Pterygota), derived from ancestors that evolved to fold their wings (Neoptera) and possess biting mouthparts, but undergo little or no metamorphosis.


Protodiptera is an extinct order of insects containing the two genera Permotipula and Permila.


Psocodea is a taxonomic group of insects comprising the bark lice, book lice and true lice. It was formerly considered a superorder, but is now generally considered by entomologists as an order. Despite the greatly differing appearance of lice, they are believed to have evolved from within the former order "Psocoptera", which contained the bark lice and book lice. Psocodea contains around 11,000 species, divided among seven suborders.


The Pterygota are a subclass of insects that includes the winged insects. It also includes insect orders that are secondarily wingless (that is, insect groups whose ancestors once had wings but that have lost them as a result of subsequent evolution).The pterygotan group comprises almost all insects. The insect orders not included are the Archaeognatha (jumping bristletails) and the Zygentoma (silverfishes and firebrats), two primitively wingless insect orders. Also not included are the three orders no longer considered to be insects: Protura, Collembola, and Diplura.

Insect orders
Extant Notoptera families


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