from Wikipedia, the free encyclopedia
Mantophasma zephyrum

Mantophasma zephyrum

Trunk : Arthropod (arthropoda)
Superclass : Six-footed (Hexapoda)
Class : Insects (Insecta)
Subclass : Flying insects (Pterygota)
Superordinate : New winged wing (Neoptera)
Order : Gladiators
Scientific name
Zompro , Klass , Kristensen & Adis , 2002

The gladiators or gladiator horrors (Mantophasmatodea), sometimes also called heel walkers , are an order of carnivorous insects that was first described in 2002 .


The name Mantophasmatodea is a combination of the scientific names of the praying mantis (Mantodea) and the ghosts (Phasmatodea), since the gladiators appear at first glance to be a combination of both orders. The name Gladiators is inspired by the author of the order, Oliver Zompro . He compared the helmet armor of a gladiator in the film of the same name with the conspicuous bristling of the head of an animal later described as a tyrannophasma gladiator , which he examined while watching the film on a DVD . Since the last link of the feet, the arolium , is almost always held up, the representatives of this order are also called heel runners.


All gladiators are wingless and have a downward-pointing head with long antennae , which in adults are divided into a 14-membered basiflagellum and a 7-membered distiflagellum. The three breast segments overlap with the rear edge in each case on the following segment. The front and middle legs are thickened and armed with thorns on the inside. The feet ( tarsi ) are five-limbed, the third limb has a triangular extension on the top. Between the claws there is the relatively large arolium on which a bristle ring sits. The abdomen is made up of ten segments. The enlarged and asymmetrical cerci in the male consist of one segment. Depending on the species, the animals reach a length of 1.5 to 4 cm, with the females always being larger than the males.

Occurrence and way of life

Larva of a gladiator

All species still alive today are found in southern Africa , more precisely in Namibia , Tanzania and South Africa . They feed on small insects, which they hold on to with their front legs and - depending on the size of the prey - also with their middle legs. For intra-species communication and here primarily to find a partner, they drum their abdomen on the floor and thus generate species- and gender-specific substrate vibrations. The female feels the vibrations generated by the male and responds to them by also starting to drum. The male follows this drumming and thus finds the way to the female. To begin mating , which lasts several hours to three days , the male jumps on the female. After the sperm has been transferred, the male gets to safety from the female by jumping as long as possible (up to ten centimeters). The elongated eggs, which do not have a removable lid, are glued together with secretion and sand to form an "egg pod" and deposited in the bottom. Due to the plastron- like egg surface, the eggs can survive for some time if they are flooded. The eggs survive the dry summer, only in the next rainy season the nymphs hatch and moult five times within a few weeks. The arolium , which is normally stretched away from the ground, is lowered to the ground in the event of danger, strong wind or rain and thus gives the insects a better grip. The gladiators are hemimetabolic insects .


The order was only drawn up in 2002 by Oliver Zompro, Joachim Adis (both Max Planck Institute for Limnology in Plön ), Klaus-Dieter Klass ( Senckenberg Natural History Collections Dresden ) and Niels-Peder Kristensen ( Copenhagen ), making it the first newly described insect order since 1914. Some specimens that had previously existed in museum collections and could not be identified were not recognized as being of this kind. In 1999 Zompro found 45 million year old insects in Baltic amber († Raptophasma kerneggeri ) while collecting material for his dissertation on the systematics of ghost horrors , and during further research in 2000 and 2001, the aforementioned museum specimens ( Mantophasma zephyrum and Mantophasma subsolanum ). In summer 2001 the biologist Martin Wittneben from the University of Bremen carried out research on the Brandberg massif (2600 m) in Namibia for his diploma thesis on the vegetation of this island mountain range in the middle of the Namib desert , accompanied by his Swiss colleague Hansueli Dubach. During this expedition, the researchers came across an extraordinary insect that Wittneben brought to Bremen. In investigations at the University of Bremen with the participation of the biologist Dr. Hartmut Koehler found that this could not be assigned to any known insect order. It was then made available to the curator of the collection of the National Museum in Windhoek , Eugène Marais. At that time, Adis and Zompro made a number of inquiries regarding the found museum specimens to African museums, including the one in Windhoek. Marais recognized the animal Wittneben had found on the pictures he sent. Thanks to Wittneben's precise information on the location of his individual, an excursion was carried out in 2002 on the Brandberg and in the Erongo Mountains in Namibia by Zompro, Marais, John Irisch (from the Namibian National Biodiversity Program in Windhoek) and others, on which further, living representatives were found .


External system

The relationship of the Mantophasmatodea to other insect orders has not yet been clearly clarified. This order of insects is placed either in the vicinity of the praying mantis (Mantodea), the cricket cockroaches (Grylloblattodea), or the ghosts (Phasmatodea).
Some authors see in the gladiators only a subordination of the order Notoptera , to which then the cricket cockroaches (Grylloblattodea) are counted as a sister group and thus second subordination.
Other authors see in the Mantophasmatodea an independent order within the superordinate order of the Neuflügler (Neoptera), more precisely the Polyneoptera, a systematically as yet unexplained group. From a phylogenetic perspective, stick insects (Phasmatodea) that are in this the Orthopteromorpha to which the gladiators are counted, distinguished from the Phasmatomorpha to which, among other embioptera (Embioptera) and stoneflies (Plecoptera) are counted:


 Cricket cockroaches (Grylloblattodea or Notoptera)


 Earwigs (Dermaptera)


 Mantis (Mantodea)


 Cockroaches (Blattodea), termites (Isoptera) and ground lice (Zoraptera)


 Gladiator horror (Mantophasmatodea)


 Long- probe horror (Ensifera)


 Short-antennae terrors (Caelifera)

Template: Klade / Maintenance / 3

Template: Klade / Maintenance / 3

Alternatively, the following system can be found, which divides the new winged birds (Neoptera) into paurometabola and eumetabola. According to this classification, the gladiators also belong to the Paurometabola and thus to the closer relationship of the ghost horrors. The question marks, "?", Mark a currently discussed and controversial position of the taxon:


 Stoneflies (Plecoptera)


 Embioptera (Embioptera)


 Cricket cockroaches (Grylloblattodea or Notoptera)


 Earwigs (Dermaptera)


 Mantis (Mantodea)




 ? Gladiator horror (Mantophasmatodea)


 ? Long- probe horror (Ensifera)


 ? Short-antennae terrors (Caelifera)


 ? Ghost horror (Phasmatodea)

Template: Klade / Maintenance / 3Template: Klade / Maintenance / 4

Internal system

The order was originally described with only one family , the Mantophasmatidae . At first only the two species of the recent genus Mantophasma and the already mentioned species of the extinct genus Raptophasma belonged to it . Meanwhile, much more fossil and recent species are differentiated.

The latest family tree analysis using mitochondrial genes ( CO1 and 16SrRNA ) was developed by Jakob Damgaard and co-workers. In his analysis there are a total of 20 recent species, 4 of which have not yet been described (marked with sp.n.). Mantophasmatodea are divided into 3 (or 4) families with a total of 12 genera ; 3 species already described are listed as Mantophasmatodea incertae sedis (i.e. with an unrelated assignment), as precise studies of the end of the abdomen and the DNA are still lacking. In addition, following a hint from O. Zompro, the endings of the species names were changed from feminine to neuter endings due to object equality, since the word "Phasma", which is found in all generic names, is neuter (e.g. Karoophasma biedouwensis changed to K. biedouwense ).

Family Austrophasmatidae Klass , Picker , Damgaard , VanNoort , Tojo , 2003

there are also two not yet described species (Austrophasmatidae sp.n.)

Family Mantophasmatidae Zompro , Klass , Kristensen & Adis , 2002

Family Tanzaniophasmatidae Klass et al., 2003

Tyranno- / Praedatophasma group - still undescribed family

Species that have not yet been clearly assigned

Belonging to Mantophasmatodea not yet clarified:

An alternative classification, which is not based on molecular data or morphological studies of the genital apparatus, follows Zompro (2008), whereby extinct taxa , i.e. their species only exist as fossils with "†" and those with not yet established (generic) assignment "?" Marked are:

  • † Ensiferophasmatidae Zompro , 2005
    • † Ensiferophasmatinae Zompro , 2005
      • † Ensiferophasmatini Zompro , 2005
        • Ensiferophasma Zompro , 2005
          • Ensiferophasma velociraptor Zompro , 2005
  • Mantophasmatidae Zompro , Klass , Kristensen & Adis , 2002
    • † Raptophasmatinae Zompro , 2005
      • † Raptophasmatini Zompro , 2005
    • Mantophasmatinae Zompro , Klass , Kristensen & Adis , 2002
      • Mantophasmatini Zompro , Klass , Kristensen & Adis , 2002
        (Syn. = Austrophasmatidae Klass et al. , 2003)
        • Mantophasma Zompro , Klass , Kristensen & Adis , 2002
          (Syn. = Tanzaniophasmatidae Klass et al., 2003)
          • ? Mantophasma caledonensis ( Klass et al., 2003)
            (Syn. = Austrophasma caledonensis Klass et al., 2003)
          • Mantophasma gamsbergense Zompro & Adis , 2006
          • ? Mantophasma gansbaaiensis ( Klass et al., 2003)
            (Syn. = Austrophasma gansbaaiensis Klass et al., 2003)
          • Mantophasma kudubergense Zompro & Adis , 2006
          • Mantophasma omatakoense Zompro & Adis , 2006
          • Mantophasma paresisensis ( Klass et al., 2003)
            (Syn. = Sclerophasma paresisensis Klass et al., 2003)
          • ? Mantophasma rawsonvillensis ( Klass et al., 2003)
            (Syn. = Austrophasma rawsonvillensis Klass et al., 2003)
          • Mantophasma subsolana Zompro , Klass , Kristensen & Adis , 2002
            (Syn. = Tanzaniophasma subsolana Zompro , Klass , Kristensen & Adis , 2002)
          • Mantophasma zephyra Zompro , Klass , Kristensen & Adis , 2002
      • Tyrannophasmatini Zompro , 2005
        • Tyrannophasma Zompro , 2003
          • Tyrannophasma gladiator Zompro , 2003
        • Praedatophasma Zompro & Adis , 2002
          • Praedatophasma maraisi Zompro & Adis , 2002

Individual evidence

  1. a b c Oliver Zompro: Mantophasmatodea - Gladiators in the insect kingdom. In: Arthropoda. 16 (1), March 2008, Sungaya-Verlag Kiel. ISSN  0943-7274
  2. ^ Online newspaper of the University of Vienna
  3. a b D. Hockman, MD Picker, KD Klass, L. Pretorius: Postembryonic development of the unique antenna of Mantophasmatodea (Insecta). In: Arthropod Structure and Development. 38, 2009, pp. 125-133.
  4. MJB Eberhard, MD Picker: Vibrational Communication in two sympatric species of Mantophasmatodea (Heelwalkers). In: Journal of Insect Behavior. 21, 2008, pp. 240-257.
  5. APA press release
  6. MJB Eberhard, G. Pass, MD Picker, R. Beutel, R. Predel, SN Gorb: Structure and Function of the Arolium of Mantophasmatodea (Insecta). In: Journal of Morphology. 270, 2009, pp. 1247-1261.
  7. Klaus-Dieter Klass, Oliver Zompro, Niels-Peder Kristensen, Joachim Adis: Mantophasmatodea: A New Insect Order with Extant Members in the Afrotropics. In: Science. 296, 2002, pp. 1456-1459.
  8. Oliver Zompro: A generic revision of the insect order Phasmatodea: Areolatae, including a new order of insects. Max Planck Institute for Limnology, (No. 2278), Plön 2003. (858 pages)
  9. Sensational insect discovery: Bremen biologist discovers living "fossils" in Namibia. ( Memento of June 7, 2007 in the Internet Archive ) Press release No. 219 / October 10, 2002 of the University of Bremen.
  10. R. Dallai, F. Frati, P. Lupetti, J. Adis: Sperm ultrastructure of Mantophasma zephyra (Insecta, Mantophasmatodea). In: Zoomorphology. 122, 2003, pp. 67-76.
  11. ^ MD Terry, MF Whiting: Mantophasmatodea and phylogeny of the lower neopterous Insects. In: Cladistics. 21, 2005, pp. 240-257.
  12. Cameron SL, Barker SC, Whiting MF: Mitochondrial genomics and the new insect order Mantophasmatodea. In: Molecular Phylogenetics and Evolution. 38, 2006, pp. 274-279.
  13. Antonio Arillo, Michael S. Engel: rock crawlers in Baltic amber (notoptera, Mantophasmatodea). (= American Museum novitates . No. 3539). New York 2006. ( online version )
  14. Oliver Zompro: Inter- and Intraordinal relationship of the Mantophasmatodea, with comments on the phylogeny of polyneopteran orders (Insecta: Polyneoptera). In: Communications from the Geological-Paleontological Institute of the University of Hamburg. 89, 2005, pp. 85-114.
  15. Jump up ↑ J. Damgaard, KD Klass, MD Picker, G. Buder: Phylogeny of the Heelwalkers (Insecta: Mantophasmatodea) based on mtDNA sequences, with evidence for additional taxa in South Africa. In: Molecular Phylogenetics and Evolution. 47 (2), 2009, pp. 443-462.
  16. MJB Eberhard, MD Picker, K.-D Klass: Sympatry in Mantophasmatodea, with the description of a new species and phylogenetic considerations. In: Organisms Diversity and Evolution. 11, 2011, pp. 43-59.

Web links

Commons : Gladiators (Mantophasmatodea)  - Collection of images, videos and audio files