Nannochoristidae

Nannochoristidae; Temporal range:
	Nannochorista philpotti
Scientific classification
Domain:	Eukaryota
Kingdom:	Animalia
Phylum:	Arthropoda
Class:	Insecta
Order:	Mecoptera
Family:	Nannochoristidae; Tillyard, 1917
Genera
	See text

Nannochoristidae is a family of scorpionflies with many unusual traits. It is a tiny, relict family with a single extant genus, Nannochorista, with eight species occurring in New Zealand, southeastern Australia, Tasmania, Argentina and Chile. Due to the group's distinctiveness from other scorpionflies, it is sometimes placed in its own order, the Nannomecoptera. Some studies have placed them as the closest living relatives of fleas.[1] Most mecopteran larvae are eruciform, or shaped like caterpillars. Nannochoristid larvae, however, are elateriform, and have elongated and slender bodies. The larvae are aquatic, which is unique among mecopterans. The larvae are predatory, hunting on the beds of shallow streams, primarily on the larvae of aquatic Diptera like chironomids.[2]

Fossils indicate that Nannochoristidae formerly had a wider distribution, being present in the Northern Hemisphere during the Jurassic and Early Cretaceous.[3]

Some research suggests the nannochoristids are the only holometabolous insects with true larval compound eyes.[4] All other eyed larvae have stemmata, which are structurally different from adult compound eyes with ommatidia. This is unusual, since most adult features are present as imaginal discs in larvae and not formed until pupation. The presence of compound eyes in nannochoristid larvae suggests the timing of the development of adult features can be initiated earlier in development, which has important implications for insect evolutionary development.

Phylogeny

The cladogram of external relationships, based on a 2008 DNA and protein analysis, shows the family as a clade, sister to the Siphonaptera (fleas) and rest of Mecoptera, and more distantly related to the Diptera (true flies) and Mecoptera (scorpionflies).[5][6][7][8]

Panzygothoraca

Panorpida

Antliophora

Diptera (true flies)

Mecopteroidea

Nannochoristidae

Mecoptera (scorpionflies, hangingflies) (exc. Boreidae & Nannochoristidae)

	Boreidae (snow scorpionflies)

	Siphonaptera (fleas)

Amphiesmenoptera

	Trichoptera (caddisflies)

	Lepidoptera (butterflies and moths)

Hymenoptera (sawflies, wasps, ants, bees)

(part of Endopterygota)

A more recent study in 2021 found Nannochoristidae to be the sister group to fleas, with strong support.[1]

Antliophora

Diptera (true flies)

Boreidae (snow scorpionflies, 30 spp.)

	Nannochoristidae (southern scorpionflies, 8 spp.)

	Siphonaptera (fleas, 2500 spp.)

Pistillifera (scorpionflies, hangingflies, 400 spp.)

Genera

After Cao et al. 2022.[9]

Nannochorista Tillyard, 1917 (8 species: Argentina, Chile, Tasmania, Australia, New Zealand) Koonwarra fossil bed, Australia, Early Cretaceous (Aptian)
†Dahurochorista Sukatsheva 1985 (one species: Ichetuy Formation, Russia, Late Jurassic (Oxfordian)
†Dahurolarva Sukatsheva 1985 (one species: Ichetuy Formation, Oxfordian, Russia) (based on larvae only)
†Itaphlebia Sukatsheva 1985 (14 species) Itat Formation, Russia, Middle Jurassic (Bathonian), Daohugou, China, Middle Jurassic (Callovian), Karabastau Formation, Kazakhstan, Late Jurassic (Oxfordian) Doronino Formation, Russia, Early Cretaceous (Barremian)
†Namdyrus Sukacheva 1993 Khaya Formation, Russia, Late Jurassic (Tithonian)
†Tarantogus Sukatsheva, 1985 Itat Formation, Russia, Bathonian (based on larvae only)
†Undisca Sukatsheva, 1990 Glushkovo Formation, Russia, Tithonian

References

Tihelka, Erik; Giacomelli, Mattia; Huang, Di-Ying; Pisani, Davide; Donoghue, Philip C. J.; Cai, Chen-Yang (2020-12-21). "Fleas are parasitic scorpionflies". Palaeoentomology. 3 (6): 641–653–641–653. doi:10.11646/palaeoentomology.3.6.16. hdl:1983/8d3c12c6-529c-4754-b59d-3abf88a32fc9. ISSN 2624-2834. S2CID 234423213.
Fraulob, Maximilian; Wipfler, Benjamin; Hünefeld, Frank; Pohl, Hans; Beutel, Rolf G. (March 2012). "The larval abdomen of the enigmatic Nannochoristidae (Mecoptera, Insecta)". Arthropod Structure & Development. 41 (2): 187–198. doi:10.1016/j.asd.2011.11.001. PMID 22309703.
LIU, NAN; Zhao, Yunyun; REN, DONG (2010-04-07). "Two new fossil species of Itaphlebia (Mecoptera: Nannochoristidae) from Jiulongshan Formation, Inner Mongolia, China". Zootaxa. 2420 (1): 37. doi:10.11646/zootaxa.2420.1.3. ISSN 1175-5334.
Melzer, R. R.; H. F. Paulus & N. P. Kristensen (1994). "The larval eye of nannochoristid scorpionflies (Insecta, Mecoptera)". Acta Zoologica. 75 (3): 201–208. doi:10.1111/j.1463-6395.1994.tb01207.x.
Whiting, Michael F.; Whiting, Alison S.; Hastriter, Michael W.; Dittmar, Katharina (2008). "A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations". Cladistics. 24 (5): 677–707. CiteSeerX 10.1.1.731.5211. doi:10.1111/j.1096-0031.2008.00211.x. S2CID 33808144.
Yeates, David K.; Wiegmann, Brian. "Endopterygota Insects with complete metamorphosis". Tree of Life. Retrieved 24 May 2016.
Whiting, Michael F. (2002). "Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera". Zoologica Scripta. 31 (1): 93–104. doi:10.1046/j.0300-3256.2001.00095.x. S2CID 56100681. Archived from the original on 2013-01-05.
Wiegmann, Brian; Yeates, David K. (2012). The Evolutionary Biology of Flies. Columbia University Press. p. 5. ISBN 978-0-231-50170-5.
Cao, Yizi; Lin, Xiaodan; Shih, Chungkun; Ren, Dong (2022-06-24). "Two new species of Itaphlebia (Insecta, Mecoptera, Nannochoristidae) from the late Middle Jurassic of China". ZooKeys (1108): 175–188. doi:10.3897/zookeys.1108.85378. ISSN 1313-2970. PMC 9848664. PMID 36760696.

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[:0-1] Tihelka, Erik; Giacomelli, Mattia; Huang, Di-Ying; Pisani, Davide; Donoghue, Philip C. J.; Cai, Chen-Yang (2020-12-21). "Fleas are parasitic scorpionflies". Palaeoentomology. 3 (6): 641–653–641–653. doi:10.11646/palaeoentomology.3.6.16. hdl:1983/8d3c12c6-529c-4754-b59d-3abf88a32fc9. ISSN 2624-2834. S2CID 234423213.

[2] Fraulob, Maximilian; Wipfler, Benjamin; Hünefeld, Frank; Pohl, Hans; Beutel, Rolf G. (March 2012). "The larval abdomen of the enigmatic Nannochoristidae (Mecoptera, Insecta)". Arthropod Structure & Development. 41 (2): 187–198. doi:10.1016/j.asd.2011.11.001. PMID 22309703.

[3] LIU, NAN; Zhao, Yunyun; REN, DONG (2010-04-07). "Two new fossil species of Itaphlebia (Mecoptera: Nannochoristidae) from Jiulongshan Formation, Inner Mongolia, China". Zootaxa. 2420 (1): 37. doi:10.11646/zootaxa.2420.1.3. ISSN 1175-5334.

[Melzer1994-4] Melzer, R. R.; H. F. Paulus & N. P. Kristensen (1994). "The larval eye of nannochoristid scorpionflies (Insecta, Mecoptera)". Acta Zoologica. 75 (3): 201–208. doi:10.1111/j.1463-6395.1994.tb01207.x.

[Whiting2008-5] Whiting, Michael F.; Whiting, Alison S.; Hastriter, Michael W.; Dittmar, Katharina (2008). "A molecular phylogeny of fleas (Insecta: Siphonaptera): origins and host associations". Cladistics. 24 (5): 677–707. CiteSeerX 10.1.1.731.5211. doi:10.1111/j.1096-0031.2008.00211.x. S2CID 33808144.

[6] Yeates, David K.; Wiegmann, Brian. "Endopterygota Insects with complete metamorphosis". Tree of Life. Retrieved 24 May 2016.

[Whiting2002-7] Whiting, Michael F. (2002). "Mecoptera is paraphyletic: multiple genes and phylogeny of Mecoptera and Siphonaptera". Zoologica Scripta. 31 (1): 93–104. doi:10.1046/j.0300-3256.2001.00095.x. S2CID 56100681. Archived from the original on 2013-01-05.

[WiegmannYates2012-8] Wiegmann, Brian; Yeates, David K. (2012). The Evolutionary Biology of Flies. Columbia University Press. p. 5. ISBN 978-0-231-50170-5.

[9] Cao, Yizi; Lin, Xiaodan; Shih, Chungkun; Ren, Dong (2022-06-24). "Two new species of Itaphlebia (Insecta, Mecoptera, Nannochoristidae) from the late Middle Jurassic of China". ZooKeys (1108): 175–188. doi:10.3897/zookeys.1108.85378. ISSN 1313-2970. PMC 9848664. PMID 36760696.