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
    |--Mantophasmatodea TW05
    |    |  i. s.: Adicophasma spinosa GE05
    |    |         Raptophasma Zompro 2001 KP03
    |    |           `--R. kerneggeri Zompro 2001 KZ02
    |    |         Praedatophasma maraisi Zompro & Adis 2002 KP03
    |    |--Tanzaniophasma Klass, Picker et al. 2003 [Tanzaniophasmatidae] KP03
    |    |    `--*T. subsolana (Zompro, Klass et al. 2002) [=Mantophasma subsolana] KP03
    |    |--Mantophasmatidae TW05
    |    |    |--Tyrannophasma gladiator TW05
    |    |    |--Sclerophasma Klass, Picker et al. 2003 KP03
    |    |    |    `--*S. paresisensis Klass, Picker et al. 2003 KP03
    |    |    `--Mantophasma Zompro, Klass et al. 2002 KP03
    |    |         `--*M. zephyra Zompro, Klass et al. 2002 KP03
    |    `--Austrophasmatidae KP03
    |         |  i. s.: Namaquaphasma Klass, Picker et al. 2003 KP03
    |         |           `--*N. ookiepensis Klass, Picker et al. 2003 KP03
    |         |--+--Hemilobophasma Klass, Picker et al. 2003 KP03
    |         |  |    `--*H. montaguensis Klass, Picker et al. 2003 KP03
    |         |  `--Karoophasma Klass, Picker et al. 2003 KP03
    |         |       |--*K. biedouwensis Klass, Picker et al. 2003 KP03
    |         |       `--K. botterkloofensis Klass, Picker et al. 2003 KP03
    |         `--+--Lobophasma Klass, Picker et al. 2003 KP03
    |            |    `--*L. redelinghuysensis Klass, Picker et al. 2003 KP03
    |            `--Austrophasma Klass, Picker et al. 2003 KP03
    |                 |--*A. caledonensis Klass, Picker et al. 2003 KP03
    |                 |--A. gansbaaiensis Klass, Picker et al. 2003 KP03
    |                 `--A. rawsonvillensis Klass, Picker et al. 2003 KP03
    `--Grylloblattodea [Grylloblattaria, Grylloblattoidea, Notoptera] TW05
         |  i. s.: Olgaephilus St02
         |         Haplopterum St02
         |         Kargarella St02
         |         Thaumatophora pronotalis Si02
         |         Sylvonympha tshekardensis St02
         |--Protoperlina GE05
         `--+--Nele Ansorge 1996 BN05 [Neleidae St02]
            |--Kortshakoliidae St02
            |--Mesorthopteron F71 [Mesorthopteridae St02]
            |    `--M. locustoides F71
            |--Archiprobnis BN02 [Archiprobnidae St02, Archiprobnisidae]
            |    |--A. conjuncta BN02
            |    `--A. repens BN02
            |--Geinitziidae [Prosepididontidae] St02
            |    |--Prosepididontes calopterix St02
            |    |--Shurabia St02
            |    `--Fletchitzia picturata RJ93
            `--+--Ideliidae St02
               |    |--Ideliopsina nana Z02
               |    |--Sojanidelia floralis St02
               |    |--Archidelia ovata St02
               |    |--Sylvidelia latipennis St02
               |    `--Austroidelia perplexa Riek 1954 K-P91, F71
               `--+--Permotermopsidae St02
                  `--+--Megakhosarodes RJ93 [Megakhosaridae St02]
                     |    `--M. zajsanicus RJ93
                     `--+--Blattogryllidae St02
                        |    |--Dorniella St02
                        |    |--Blattogryllus karatavicus GE05
                        |    |--Protoblattogryllus zajsanicus Storozhenko 1990 RJ93
                        |    `--Parablattogryllus obscurus Storozhenko 1988 RJ93
                        `--Grylloblattidae St02
                             |--Grylloblattella GE05
                             |--Grylloblattina djakonovi GE05, RD77
                             |--Namkungia GE05
                             |--Galloisiana G61
                             |    |--G. chujoi Gurney 1961 G61
                             |    |--G. kiyosawai Asahina 1959 G61
                             |    |--G. nipponensis G61
                             |    |--G. notabilis Silvestri 1927 I92
                             |    `--G. yuasai Asahina 1959 G61
                             `--Grylloblatta TW05
                                  |--G. barberi G61
                                  |--G. bifratrilecta G61
                                  |--G. campodeiformis Walker 1914 KP03
                                  |    |--G. c. campodeiformis R00
                                  |    `--G. c. occidentalis R00
                                  |--G. chirurgica Gurney 1961 G61
                                  |--G. rothi G61
                                  |--G. sculleni G61
                                  `--G. washoa Gurney 1961 G61

*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 10^th 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.