Diagram of holotype of the Late Carboniferous Parahomalophlebia courtini, from Rasnitsyn (2002c).

Belongs within: Pterygota.
Contains: Forficulida, Embiidina, Protoperlaria, Plecoptera, Archaeorthoptera, Strephocladina, Dictyoptera, Holophasmatodea, Xenonomia.

The Polyneoptera are a major subdivision of the insects, including grasshoppers, cockroaches and related taxa. Though members of this group are often broadly similar in habitus, its monophyly remains uncertain.

The name Perlidea was used in Rasnitsyn & Quicke (2002) for a group including the Plecoptera, Grylloblattodea, Dermaptera and Embioptera. Molecular analysis (Terry & Whiting 2005) supports a clade including most of these taxa (excluding the Embioptera), but with the Dictyoptera and Zoraptera also nested within it.

The Orthopteroidea is an insect clade supported by molecular analysis (Terry & Whiting 2005), including the grasshoppers, stick insects and webspinners.

A choir of Zoraptera
Zorapteran, from the Papua Insects Foundation.

The Zoraptera are perhaps the most obscure of all insect orders. Zorapterans are inhabitants of rotting logs in tropical forests. A few species range north of the tropics in North America and Asia. Until recently, they were regarded as quite rare but apparently they have turned out to be not uncommon in suitable habitat. Zoraptera are semi-social, living in colonies of up to a hundred or so individuals.

The name “Zoraptera” means “purely wingless” (the reason for the name should be obvious), and an alternative translation of “zoros” (the same element as in the beginning of “Zoroaster”, I believe) explains the occassionally used common name for Zoraptera of “angel insects”. There is absolutely no rational justification for calling Zoraptera angels, but the name is just poetic enough that I hope it catches on. It is now known that winged Zoraptera do exist, just not very often. Because rotting logs at just the right stage of decomposition are a temporary resource, zorapterans have an aphid-like life cycle, with blind, wingless individuals breeding and multiplying in their log until resources start running out, at which time winged individuals with eyes start to emerge. These winged individuals are able to leave the doomed colony and seek out a new piece of suitable habitat elsewhere. Notably, the majority of winged dispersers are female—males are quite rare. Female dispersers probably mate with males of their parent colony before dispersing.

Most authors include living species in a single genus, Zorotypus. Kukalová-Peck & Peck (1993) did establish a number of new genera, but as their classification was based on wing characters and could only be applied to a selection of recent taxa (leaving those species for which winged individuals were unknown in a Zorotypus of convenience) it has not been widely accepted. A species from Cretaceous amber has been placed in a distinct genus, Xenozorotypus, with species of the modern genus Zorotypus also known from the same time period (Engel & Grimaldi 2002).

The phylogenetic relationships of Zoraptera are rather obscure, to say the least. To quote Engel & Grimaldi (2002): “At one time or another Zoraptera has been considered sister to Isoptera (Boudreaux, 1979; Caudell, 1918; Crampton, 1920; Weidner, 1969, 1970), Isoptera + Blattaria (Silvestri, 1913), Paraneoptera (Hennig, 1953, 1969, 1981; Kristensen, 1975), Embiidina (Minet and Bourgoin, 1986; Engel and Grimaldi, 2000; Grimaldi, 2001), Holometabola (Rasnitsyn, 1998), Dermaptera (Carpenter and Wheeler, 1999), Dermaptera + Dictyoptera (Kukalová-Peck and Peck, 1993); basal within Thysanoptera (Karny, 1922) or Psocoptera (Karny, 1932); or unresolved within either basal Neoptera (Kristensen, 1991) or Orthoptera, Phasmida, and Embiidina (Kukalová-Peck, 1991).” At present it is pretty well-accepted that Zoraptera are somewhere within the Polyneoptera, the clade or grade including the cockroaches, crickets, etc., but getting more resolution than this is still difficult. Engel & Grimaldi (2002) favour a relationship to Embioptera, while Terry & Whiting (2005) link them to Dermaptera (earwigs). Rasnitsyn (2002b) points out that the characters Engel & Grimaldi used to link Zoraptera to Embioptera are prone to homoplasy, but his own suggestion of a sister-relationship to Holometabola is poorly supported.

Systematics of Polyneoptera

Characters (from Rasnitsyn 2002c, as Gryllones): Large to medium size, rarely small. Mouthparts chewing. Unless lost, wings resting flat over body (neither roof-like nor spread), with contralateral wings overlapping each other in part or completely, with hind wing anal area usually enlarged and, when folded, bent down along line running before 2A (occasionally before 1A), with common base of anal veins reverted upside down. Thoracic venter usually not cryptosternous. Third valve (gonocoxite 9 with or without its style) forming main intromittent part of ovipositor. Male gonostylus free or, usually, lost. Paracercus lost. Ovaries usually panoistic. Development embryonicised, eclosing nymph with compound eyes, ocelli and usually with full number of tarsomeres, and usually without adult moulting.

<==Polyneoptera (see below for synonymy)
    |--Thoronysis [Thoronysidae]R02c
    |--Prototettix [Prototettigidae]R02c
    |    `--P. lobulatusR15
    |--Eucaenus [Eucaenidae]R02c
    |    `--E. ovalis Scudder 1885BN02
    |    |--PropteticusR02c
    |    |--DieconeuraR02c
    |    `--Axiologus Handlirsch 1906R02c, BN02
    |         `--A. thoracicus Handlirsch 1906BN02
    `--+--Parahomalophlebia [Homalophlebiidae]R02c
       |    `--P. courtiniR02c
          |  |  `--+--EmbiidinaBB16
          |  |     `--Brachyphyllophagus [Brachyphyllophagidae]R02d
          |  |          `--B. phasmaR02d
          |  `--Zoraptera [Zorotypida]TW05
          |       |--Xenozorotypus burmiticusGE05
          |       `--Zorotypus Silvestri 1913BB16, G38 [Zorotypidae]
          |            |--*Z. guineensis Silvestri 1913G38
          |            |--Z. barberi Gurney 1938G38
          |            |--Z. buxtoni Karny 1932G38
          |            |--Z. caudelli Karney 1926G38
          |            |--Z. ceylonicus Silvestri 1913G38
          |            |--Z. cramptoni Gurney 1938G38
          |            |--Z. goeletiGE05
          |            |--Z. hubbardi Caudell 1918G38
          |            |--Z. javanicus Silvestri 1913G38
          |            |--Z. longicercatus Caudell 1927G38
          |            |--Z. manni Caudell 1923 [=Zoraptera (l. c.) manni]G38
          |            |--Z. nascimbeneiGE05
          |            |--Z. neotropicus Silvestri 1916G38
          |            |--Z. palaeus Poinar 1988RJ93
          |            |--Z. philippinensis Gurney 1938G38
          |            |--Z. shannoni Gurney 1938G38
          |            |--Z. silvestrii Karny 1926G38
          |            |--Z. snyderi Caudell 1920G38
          |            `--Z. swezeyi Caudell 1922G38
             |  `--+--PlecopteraBB16
             |     `--Probnis Sellards 1909BNL05 [ProbnidaeGE05]
                   |  `--XenonomiaBB16
                   `--+--Blattinopseida [Blattinopseidae, Blattinopsodea]BN02
                      |    |--BlattinopsisR02b
                      |    `--GlaphyrophlebiaR02b
                      |         |--G. subcostalisR02b
                      |         `--G. uralensisR02b
                      `--Paoliida [Protoptera]NR13
                           |--Stygne [Stygneidae]RZ02
                           |    `--S. roemeriR02a
                                |--Kemperala Brauckmann 1984IB11
                                |    `--K. hagenensis Brauckmann 1984IB11
                                |    |--Z. grandis Kukalová 1958K-P83
                                |    `--Z. occidentalis Laurentiaux-Vieira & Laurentiaux 1986RJ93
                                `--Holasicia Kukalová 1958IB11
                                     |--H. rasnitsyni Bracukmann 1984IB11
                                     `--H. vetulaR02a
Polyneoptera incertae sedis:
  Chresmodidae [Chresmododea]R02c
    |--Gryllidium oweniR02c
    `--Chresmoda Germar 1839 [incl. Propygolampis Weyenbergh 1874]R02c
         |--C. aquatica Martinez-Delclòs 1989RJ93
         |--C. obscura [incl. Propygolampis giganteus]R02c
         `--C. orientalisR02c
  Polyernus complanatusR02c
  Lodevopterum Béthoux, Nel et al. 2005BNL05
    `--*L. angustus Béthoux, Nel et al. 2005BNL05
  Protoblattina Meunier 1909 [Protoblattidae, Protoblattinidae]BNL05
    `--*P. bouvieri Meunier 1909BNL05
  Liomopterella vulgaris Sharov 1961BNL05
    |--Paraphenopterum Storozhenko 1992BNL05
    `--Laurentiaupterum Béthoux, Nel et al. 2005BNL05
         `--*L. elegantissimum (Meunier 1921) [=Polyetes elegantissima]BNL05
  Polyetes Handlirsch 1906BNL05
  Pteroferganella Gorokhov 1989BN02

Polyneoptera [Aetioptera, Anartioptera, Blattopteriformia, Cacurgida, Eoblattida, Eukinolabia, Gryllidea, Grylloblattida, Grylloblattina, Gryllones, Haplocercata, Mantides, Orthopterae, Orthopterida, Orthopterodida, Orthopteroidea, Orthopteromorpha, Paurometabola, Perlidea, Phasmatomorpha, Plecopterida, Plecopterodea, Plecopteroidea, Polyorthoptera, Protoblattodea, Ulonata]

*Type species of generic name indicated


[BB16] Bai, M., R. G. Beutel, K.-D. Klass, W. Zhang, X. Yang & B. Wipfler. 2016. Alienoptera—a new insect order in the roach-mantodean twilight zone. Gondwana Research 39: 317–326.

[BN02] Béthoux, O., & A. Nel. 2002. Venation pattern and revision of Orthoptera sensu nov. and sister groups. Phylogeny of Palaeozoic and Mesozoic Orthoptera sensu nov. Zootaxa 96: 1–88.

[BNL05] Béthoux, O., A. Nel, J. Lapeyrie & G. Gand. 2005. New data on Paleozoic grylloblattid insects (Neoptera). Journal of Paleontology 79 (1): 125–138.

Engel, M. S., & D. A. Grimaldi. 2002. The first Mesozoic Zoraptera (Insecta). American Museum Novitates 3362: 1–20.

[GE05] Grimaldi, D., & M. S. Engel. 2005. Evolution of the Insects. Cambridge University Press: New York.

[G38] Gurney, A. B. 1938. A synopsis of the order Zoraptera, with notes on the biology of Zorotypus hubbardi Caudell. Proceedings of the Entomological Society of Washington 40 (3): 57–87.

[IB11] Ilger, J.-M., & C. Brauckmann. 2011. The smallest Neoptera (Baryshnyalidae fam. n.) from Hagen-Vorhalle (early Late Cretaceous: Namurian B; Germany). ZooKeys 130: 91–102.

[K-P83] Kukalová-Peck, J. 1983. Origin of the insect wing and wing articulation from the arthropodan leg. Canadian Journal of Zoology 61: 1618–1669.

Kukalová-Peck, J., & S. B. Peck. 1993. Zoraptera wing structures: Evidence for new genera and relationship with the blattoid orders (Insecta: Blattoneoptera). Systematic Entomology 18: 333–350.

[NR13] Nel, A., P. Roques, P. Nel, A. A. Prokin, T. Bourgoin, J. Prokop, J. Szwedo, D. Azar, L. Desutter-Grandcolas, T. Wappler, R. Garrouste, D. Coty, D. Huang, M. S. Engel & A. G. Kirejtshuk. 2013. The earliest known holometabolous insects. Nature 503: 257–261.

[R02a] Rasnitsyn, A. P. 2002a. Infraclass Scarabaeones Laicharting, 1781. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 84–85. Kluwer Academic Publishers: Dordrecht.

[R02b] Rasnitsyn, A. P. 2002b. Cohors Cimiciformes Laicharting, 1781. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 104–115. Kluwer Academic Publishers: Dordrecht.

[R02c] Rasnitsyn, A. P. 2002c. Infraclass Gryllones Laicharting, 1781. The grylloneans (=Polyneoptera Martynov, 1938). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 254–262. Kluwer Academic Publishers: Dordrecht.

[R02d] Rasnitsyn, A. P. 2002d. Order Embiida Burmeister, 1835. The webspinners (=Embioptera Shipley, 1904). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 291–293. Kluwer Academic Publishers: Dordrecht.

[RZ02] Rasnitsyn, A. P., & V. V. Zherikhin. 2002. Impression fossils. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 437–444. Kluwer Academic Publishers: Dordrecht.

[R15] Rehn, J. A. G. 1915. A further contribution to the knowledge of the Orthoptera of Argentina. Proceedings of the Academy of Natural Sciences of Philadelphia 67 (2): 270–292.

[RJ93] Ross, A. J., & E. A. Jarzembowski. 1993. Arthropoda (Hexapoda; Insecta). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 363–426. Chapman & Hall: London.

[TW05] Terry, M. D., & M. F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21: 240–257.

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