Xenonomia

Austrophasma caledonensis, photographed by Simon van Noort.

Belongs within: Polyneoptera.
Contains: Protoperlina.

The Xenonomia are a clade of insects supported by molecular analysis (Terry & Whiting 2005), uniting two relictual groups of modern wingless, carnivorous insects. The Grylloblattodea (ice crawlers) are found in cool environments in north-western North America and north-eastern Asia, and are characterised by the presence of a spina on the metathoracic sternum, and an eversible median sac on the first abdominal sternum. The Mantophasmatodea (rock crawlers or gladiators) are restricted in the modern fauna to southern Africa, though representatives are also known from Baltic amber; they possess a hypognathous head, and an enlarged pretarsal arolium with a fringe of long setae. Both groups also lack ocelli (Grimaldi & Engel 2005). Male Mantophasmatodea have essentially symmetrical genitalia and strongly curved cerci in Tanzaniophasma subsolana whereas members of the families Mantophasmatidae and Austrophasmatidae have asymmetrical genitalia and less strongly curved cerci (Klass et al. 2003).

A number of winged fossil insects have also been associated with the Grylloblattodea, though it is uncertain whether these taxa form a monophyletic group. Alternatively, at least some winged ‘grylloblattodeans’ may represent stem-xenonomians (Grimaldi & Engel 2005).

<==Xenonomia
    |--MantophasmatodeaTW05
    |    |  i. s.: Adicophasma spinosaGE05
    |    |         Raptophasma Zompro 2001KP03
    |    |           `--R. kerneggeri Zompro 2001KZ02
    |    |         Praedatophasma maraisi Zompro & Adis 2002KP03
    |    |--Tanzaniophasma Klass, Picker et al. 2003 [Tanzaniophasmatidae]KP03
    |    |    `--*T. subsolana (Zompro, Klass et al. 2002) [=Mantophasma subsolana]KP03
    |    |--MantophasmatidaeTW05
    |    |    |--Tyrannophasma gladiatorTW05
    |    |    |--Sclerophasma Klass, Picker et al. 2003KP03
    |    |    |    `--*S. paresisensis Klass, Picker et al. 2003KP03
    |    |    `--Mantophasma Zompro, Klass et al. 2002KP03
    |    |         `--*M. zephyra Zompro, Klass et al. 2002KP03
    |    `--AustrophasmatidaeKP03
    |         |  i. s.: Namaquaphasma Klass, Picker et al. 2003KP03
    |         |           `--*N. ookiepensis Klass, Picker et al. 2003KP03
    |         |--+--Hemilobophasma Klass, Picker et al. 2003KP03
    |         |  |    `--*H. montaguensis Klass, Picker et al. 2003KP03
    |         |  `--Karoophasma Klass, Picker et al. 2003KP03
    |         |       |--*K. biedouwensis Klass, Picker et al. 2003KP03
    |         |       `--K. botterkloofensis Klass, Picker et al. 2003KP03
    |         `--+--Lobophasma Klass, Picker et al. 2003KP03
    |            |    `--*L. redelinghuysensis Klass, Picker et al. 2003KP03
    |            `--Austrophasma Klass, Picker et al. 2003KP03
    |                 |--*A. caledonensis Klass, Picker et al. 2003KP03
    |                 |--A. gansbaaiensis Klass, Picker et al. 2003KP03
    |                 `--A. rawsonvillensis Klass, Picker et al. 2003KP03
    `--Grylloblattodea [Grylloblattaria, Grylloblattoidea, Notoptera]TW05
         |  i. s.: OlgaephilusSt02
         |         HaplopterumSt02
         |         KargarellaSt02
         |         Thaumatophora pronotalisSi02
         |         Sylvonympha tshekardensisSt02
         |--ProtoperlinaGE05
         `--+--Nele Ansorge 1996BN05 [NeleidaeSt02]
            |--KortshakoliidaeSt02
            |--MesorthopteronF71 [MesorthopteridaeSt02]
            |    `--M. locustoidesF71
            |--ArchiprobnisBN02 [ArchiprobnidaeSt02, Archiprobnisidae]
            |    |--A. conjunctaBN02
            |    `--A. repensBN02
            |--Geinitziidae [Prosepididontidae]St02
            |    |--Prosepididontes calopterixSt02
            |    |--ShurabiaSt02
            |    `--Fletchitzia picturataRJ93
            `--+--IdeliidaeSt02
               |    |--Ideliopsina nanaZ02
               |    |--Sojanidelia floralisSt02
               |    |--Archidelia ovataSt02
               |    |--Sylvidelia latipennisSt02
               |    `--Austroidelia perplexa Riek 1954K-P91, F71
               `--+--PermotermopsidaeSt02
                  `--+--MegakhosarodesRJ93 [MegakhosaridaeSt02]
                     |    `--M. zajsanicusRJ93
                     `--+--BlattogryllidaeSt02
                        |    |--DorniellaSt02
                        |    |--Blattogryllus karatavicusGE05
                        |    |--Protoblattogryllus zajsanicus Storozhenko 1990RJ93
                        |    `--Parablattogryllus obscurus Storozhenko 1988RJ93
                        `--GrylloblattidaeSt02
                             |--GrylloblattellaGE05
                             |--Grylloblattina djakonoviGE05, RD77
                             |--NamkungiaGE05
                             |--GalloisianaG61
                             |    |--G. chujoi Gurney 1961G61
                             |    |--G. kiyosawai Asahina 1959G61
                             |    |--G. nipponensisG61
                             |    |--G. notabilis Silvestri 1927I92
                             |    `--G. yuasai Asahina 1959G61
                             `--GrylloblattaTW05
                                  |--G. barberiG61
                                  |--G. bifratrilectaG61
                                  |--G. campodeiformis Walker 1914KP03
                                  |    |--G. c. campodeiformisR00
                                  |    `--G. c. occidentalisR00
                                  |--G. chirurgica Gurney 1961G61
                                  |--G. rothiG61
                                  |--G. sculleniG61
                                  `--G. washoa Gurney 1961G61

*Type species of generic name indicated

References

[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.

[BN05] Béthoux, O., A. Nel, J. Lapeyrie & G. Gand. 2005. New data on Paleozoic grylloblattid insects (Neoptera). Journal of Paleontology 79 (1): 125–138.

[F71] Fletcher, H. O. 1971. Catalogue of type specimens of fossils in the Australian Museum, Sydney. Australian Museum Memoir 13: 1–167.

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

[G61] Gurney, A. B. 1961. Further advances in the taxonomy and distribution of the Grylloblattidae (Orthoptera). Proceedings of the Biological Society of Washington 74: 67–76.

[I92] Iwahashi, J. (ed.) 1992. Reddo Deeta Animaruzu: a pictorial of Japanese fauna facing extinction. JICC: Tokyo.

[KP03] Klass, K.-D., M. D. Picker, J. Damgaard, S. van Noort & K. Tojo. 2003. The taxonomy, genitalic morphology, and phylogenetic relationships of southern African Mantophasmatodea (Insecta). Entomologische Abhandlungen 61 (1): 3–67.

[KZ02] Klass, K.-D., O. Zompro, N. P. Kristensen & J. Adis. 2002. Mantophasmatodea: a new insect order with extant members in the Afrotropics. Science 296: 1456–1459.

[K-P91] Kukalová-Peck, J. 1991. Fossil history and the evolution of hexapod structures. In: CSIRO. The Insects of Australia: A textbook for students and research workers vol. 1 pp. 141–179. Melbourne University Press: Carlton (Victoria).

[RD77] Richards, O. W., & R. G. Davies. 1977. Imms’ General Textbook of Entomology 10th ed. vol. 2. Classification and Biology. Chapman and Hall: London.

[R00] Ross, E. S. 2000. Embia: Contributions to the biosystematics of the insect order Embiidina. Occasional Papers of the California Academy of Sciences 149: 1–53, 1–36.

[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.

[Si02] Sinitshenkova, N. D. 2002. Order Perlida Latreille, 1810. The stoneflies (=Plecoptera Burmeister, 1839). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 281–287. Kluwer Academic Publishers: Dordrecht.

[St02] Storozhenko, S. Yu. 2002. Order Grylloblattida Walker, 1914 (=Notoptera Crampton, 1915, =Grylloblattodea Brues et Melander, 1932, +Protorthoptera Handlirsch, 1906, =Paraplecoptera Martynov, 1925, +Protoperlaria Tillyard, 1928). In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 278–281. Kluwer Academic Publishers: Dordrecht.

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

[Z02] Zherikhin, V. V. 2002. Ecological history of the terrestrial insects. In: Rasnitsyn, A. P., & D. L. J. Quicke (eds) History of Insects pp. 331–388. Kluwer Academic Publishers: Dordrecht.

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