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Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Ordo: Notoptera
Subordo: Mantophasmatodea

Familia: Mantophasmatidae
Subfamiliae (2): Mantophasmatinae - Tanzaniophasmatinae
Overview of genera (19)

AustrophasmaHemilobophasma – Karoophasma – Kuboesphasma – Lobatophasma – Mantophasma – Minutophasma – Namaquaphasma – Pachyphasma – Praedatophasma – Sclerophasma – StriatophasmaTanzaniophasmaTyrannophasma – Viridiphasma – †Adicophasma – †Ensiferophasma – †Juramantophasma – †Raptophasma
Name

Mantophasmatidae Zompro, Klass, Kristensen, & Adis, 2002

Synonyms

Austrophasmatidae
Tanzaniophasmatidae
Raptophasmatidae
Ensiferophasmatidae

References

Arillo, A.; Engel, M.S. 2006: Rock crawlers in Baltic amber (Notoptera: Mantophasmatodea). American Museum Novitates, (3539): 1–10. PDF PDF
Damgaard, J.; Klass, K.-D.; Picker, M.D.; Buder, G. 2008: Phylogeny of the heelwalkers (Insecta: Mantophasmatodea) based on mtDNA sequences, with evidence for additional taxa in South Africa. Molecular phylogenetics and evolution, 47: 443–462.
Eberhard, M.J.B.; Picker, M.D.; Klass, K.-D. 2011: Sympatry in Mantophasmatodea, with the description of a new species and phylogenetic considerations. Organisms diversity & evolution, 11(1): 43–59. DOI: 10.1007/s13127-010-0037-8
Huang, D.-y.; Nel, A.; Zompro, O.; Waller, A. 2008: Mantophasmatodea now in the Jurassic. Naturwissenschaften, 95: 947–952.
Klass, K.-D., O. Zompro, N. P. Kristensen, and J. Adis. 2002. Mantophasmatodea: a new insect order with extant members in the afrotropics. Science 296: 1456–1459.
Wipfler, B., Theska, T. & Predel, R. 2018. Mantophasmatodea from the Richtersveld in South Africa with description of two new genera and species. ZooKeys 746: 137–160. DOI: 10.3897/zookeys.746.14885. Reference page.

Vernacular names
English: gladiators, heelwalkers, rock crawlers
Nordfriisk: Haielluupern
日本語: カカトアルキ目 (踵行目)

Superregnum: Eukaryota
Regnum: Animalia
Subregnum: Eumetazoa
Cladus: Bilateria
Cladus: Nephrozoa
Cladus: Protostomia
Cladus: Ecdysozoa
Cladus: Panarthropoda
Phylum: Arthropoda
Subphylum: Hexapoda
Classis: Insecta
Cladus: Dicondylia
Subclassis: Pterygota
Ordo: Notoptera
Subordo: Mantophasmatodea

Familia: Mantophasmatidae
Subfamiliae (2): Mantophasmatinae - Tanzaniophasmatinae
Overview of genera (19)

Austrophasma – Hemilobophasma – Karoophasma – Kuboesphasma – Lobatophasma – Mantophasma – Minutophasma – Namaquaphasma – Pachyphasma – Praedatophasma – Sclerophasma – Striatophasma – Tanzaniophasma – Tyrannophasma – Viridiphasma – †Adicophasma – †Ensiferophasma – †Juramantophasma – †Raptophasma
Name

Mantophasmatidae Zompro, Klass, Kristensen, & Adis, 2002

Synonyms

Austrophasmatidae
Tanzaniophasmatidae
Raptophasmatidae
Ensiferophasmatidae

References

Arillo, A.; Engel, M.S. 2006: Rock crawlers in Baltic amber (Notoptera: Mantophasmatodea). American Museum Novitates, (3539): 1–10. PDF PDF
Damgaard, J.; Klass, K.-D.; Picker, M.D.; Buder, G. 2008: Phylogeny of the heelwalkers (Insecta: Mantophasmatodea) based on mtDNA sequences, with evidence for additional taxa in South Africa. Molecular phylogenetics and evolution, 47: 443–462.
Eberhard, M.J.B.; Picker, M.D.; Klass, K.-D. 2011: Sympatry in Mantophasmatodea, with the description of a new species and phylogenetic considerations. Organisms diversity & evolution, 11(1): 43–59. DOI: 10.1007/s13127-010-0037-8
Huang, D.-y.; Nel, A.; Zompro, O.; Waller, A. 2008: Mantophasmatodea now in the Jurassic. Naturwissenschaften, 95: 947–952.
Klass, K.-D., O. Zompro, N. P. Kristensen, and J. Adis. 2002. Mantophasmatodea: a new insect order with extant members in the afrotropics. Science 296: 1456–1459.
Wipfler, B., Theska, T. & Predel, R. 2018. Mantophasmatodea from the Richtersveld in South Africa with description of two new genera and species. ZooKeys 746: 137–160. DOI: 10.3897/zookeys.746.14885. Reference page.

Vernacular names
English: gladiators, heelwalkers, rock crawlers
Nordfriisk: Haielluupern
日本語: カカトアルキ目 (踵行目)

Mantophasmatidae is a family of carnivorous wingless insects within the order Notoptera, which was discovered in Africa in 2001.[1][2] Originally, the group was regarded as an order in its own right, and named Mantophasmatodea, but, using recent evidence indicating a sister group relationship with Grylloblattidae (formerly classified in the order Grylloblattodea),[3][4] Arillo and Engel have combined the two groups into a single order, Notoptera.[5]

Overview

The most common vernacular name for this order is gladiators, although they also are called rock crawlers, heelwalkers, mantophasmids, and colloquially, mantos.[6] Their modern centre of endemism is western South Africa and Namibia (Brandberg Massif),[7] although the modern relict population of Tanzaniophasma subsolana in Tanzania and Eocene fossils suggest a wider ancient distribution.

Mantophasmatodea are wingless even as adults, making them relatively difficult to identify. They resemble a cross between praying mantids and phasmids, and molecular evidence indicates that they are most closely related to the equally enigmatic group Grylloblattodea.[3][4] Initially, the gladiators were described from old museum specimens that originally were found in Namibia (Mantophasma zephyra) and Tanzania (M. subsolana), and from a 45-million-year-old specimen of Baltic amber (Raptophasma kerneggeri).

Live specimens were found in Namibia by an international expedition in early 2002; Tyrannophasma gladiator was found on the Brandberg Massif, and Mantophasma zephyra was found on the Erongoberg Massif.[8]

Since then, a number of new genera and species have been discovered, the most recent being two new genera, Kuboesphasma and Minutophasma, each with a single species, described from Richtersveld in South Africa in 2018.[9]

Biology

Mantophasmatids are wingless carnivores. During courtship, they communicate using vibrations transmitted through the ground or substrate.[10]

Classification

The most recent classification[9] recognizes numerous genera, including fossils:
Unidentified mantophasmid species in the Zoologische Staatssammlung München

Basal and incertae sedis
Genus: †Raptophasma Zompro, 2001 (Baltic amber, Eocene)
Genus: †Adicophasma Engel & Grimaldi, 2004 (Baltic amber, Eocene)
Genus: †Juramantophasma Huang et al, 2008[11] (Daohugou Bed, China, Callovian)
Genus: ?†Ensiferophasma Zompro, 2005 (assignment to Mantophasmatodea considered dubious)
Subfamily Tanzaniophasmatinae
Genus: Tanzaniophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – Tanzania
Species Tanzaniophasma subsolana (Zompro, Klass, Kristensen, & Adis 2002)
Subfamily Mantophasmatinae
Tribe Tyrannophasmatini
Genus: Praedatophasma Zompro & Adis, 2002 – Namibia
Species Praedatophasma maraisi Zompro & Adis, 2002
Genus: Tyrannophasma Zompro, 2003 – Namibia
Species Tyrannophasma gladiator Zompro, 2003
Tribe Mantophasmatini Zompro, Klass, Kristensen, Adis, 2002 (paraphyletic?)
Genus: Mantophasma Zompro, Klass, Kristensen, Adis, 2002 – Namibia
Species Mantophasma gamsbergense Zompro & Adis, 2006
Species Mantophasma kudubergense Zompro & Adis, 2006
Species Mantophasma omatakoense Zompro & Adis, 2006
Species Mantophasma zephyra Zompro, Klass, Kristensen, & Adis 2002
Genus: Pachyphasma Wipfler, Pohl, & Predel, 2012 – Namibia
Species Pachyphasma brandbergense Wipfler, Pohl, & Predel, 2012
Genus: Sclerophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – Namibia
Species Sclerophasma paresisense Klass, Picker, Damgaard, van Noort, & Tojo 2003
Tribe Austrophasmatini Klass, Picker, Damgaard, van Noort, Tojo, 2003
Genus: Austrophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – South Africa
Species Austrophasma caledonense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Species Austrophasma gansbaaiense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Species Austrophasma rawsonvillense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Genus: Hemilobophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – South Africa
Species Hemilobophasma montaguense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Genus: Karoophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – South Africa
Species Karoophasma biedouwense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Species Karoophasma botterkloofense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Genus: Kuboesphasma Wipfler, Theska & Predel, 2018 – South Africa
Species Kuboesphasma compactum Wipfler, Theska & Predel, 2018
Genus: Lobatophasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 (formerly Lobophasma) – South Africa
Species Lobatophasma redelinghuysense Klass, Picker, Damgaard, van Noort & Tojo, 2003
Genus: Minutophasma Wipfler, Theska & Predel, 2018 – South Africa
Species Minutophasma richtersveldense Wipfler, Theska & Predel, 2018
Genus: Namaquaphasma Klass, Picker, Damgaard, van Noort, Tojo, 2003 – South Africa
Species Namaquaphasma ookiepense Klass, Picker, Damgaard, van Noort, Tojo, 2003
Genus: Striatophasma Wipfler, Pohl & Predel, 2012 – Namibia
Species Striatophasma naukluftense Wipfler, Pohl & Predel, 2012
Genus: Viridiphasma Eberhard, Picker, Klass, 2011[12] – South Africa
Species Viridiphasma clanwilliamense Eberhard, Picker, Klass, 2011

Some taxonomists assign full family status to the subfamilies and tribes, and sub-ordinal status to the family. In total, there are 21 extant species described as of 2018.[9]
See also

Oliver Zompro

References

K.-D. Klass, O. Zompro, N.P. Kristensen, J. Adis. Mantophasmatodea: a new insect order with extant members in the afrotropics Science, 296 (2002), pp. 1456–1459
Adis, J., O. Zompro, E. Moombolah-Goagoses, and E. Marais. 2002. Gladiators: A new order of insect. Scientific American 287:60-65.
Terry, M.D., and M.F. Whiting. 2005. Mantophasmatodea and phylogeny of the lower neopterous insects. Cladistics 21(3): 240–257.
S. L. Cameron, S. C. Barker & M. F. Whiting (2006). "Mitochondrial genomics and the new insect order Mantophasmatodea". Molecular Phylogenetics and Evolution. 38 (1): 274–279. doi:10.1016/j.ympev.2005.09.020. PMID 16321547.
Arillo, A. & M. Engel (2006) Rock Crawlers in Baltic Amber (Notoptera: Mantophasmatodea). American Museum Novitates 3539:1-10
"Biodiversity Explorer: Mantophasmatodea". Iziko. Retrieved 30 June 2018.
Zompro, O.; Adis, J.; Weitschat, W. (2002). "A review of the order Mantophasmatodea (Insecta)". Zoologischer Anzeiger. 241 (3): 269–279. doi:10.1078/0044-5231-00080.
Zompro, O.; Adis, J.; Bragg, P.E.; Naskrecki, P.; Meakin, K.; Wittneben, M.; Saxe, V. (2003). "A new genus and species of Mantophasmatidae (Insecta: Mantophasmatodea) from the Brandberg Massif, Namibia, with notes on behaviour". Cimbebasia. 19: 13–24.
Wipfler, B; Theska, T; Predel, R (2018). "Mantophasmatodea from the Richtersveld in South Africa with description of two new genera and species". ZooKeys (746): 137–160. doi:10.3897/zookeys.746.14885. PMC 5904538. PMID 29674900.
Randall, J. A. (2014). "Vibrational Communication: Spiders to Kangaroo Rats". Biocommunication of Animals: 103–133. doi:10.1007/978-94-007-7414-8_7. ISBN 978-94-007-7413-1.
Huang, Di-ying; Nel, André; Zompro, Oliver; Waller, Alain (2008-06-11). "Mantophasmatodea now in the Jurassic". Naturwissenschaften. 95 (10): 947–952. doi:10.1007/s00114-008-0412-x. ISSN 0028-1042. PMID 18545982. S2CID 35408984.
Eberhard, MJB, MD Picker and KD Klass. (2011). Sympatry in Mantophasmatodea, with the description of a new species and phylogenetic considerations. Organisms Diversity & Evolution 11(1): 43-59. doi:10.1007/s13127-010-0037-8

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