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The insect order Neuroptera, or net-winged insects, includes the lacewings, mantidflies, antlions, and their relatives. The order consists of some 6,000 species. Neuroptera can be grouped together with the Megaloptera and Raphidioptera in the unranked taxon Neuropterida (once known as Planipennia) including: alderflies, fishflies, dobsonflies, and snakeflies.
Neuropterans first appeared during the Permian period, and continued to diversify through the Mesozoic era. During this time, several unusually large forms evolved, especially in the extinct family Kalligrammatidae, often called "the butterflies of the Jurassic" for their large, patterned wings.
Anatomy and biology
Neuropterans are soft-bodied insects with relatively few specialised features. They have large lateral compound eyes, and may or may not also have ocelli. Their mouthparts have strong mandibles suitable for chewing, and lack the various adaptations found in most other endopterygote insect groups.
They have four wings, usually similar in size and shape, and a generalised pattern of veins. Some neuropterans have specialised sense organs in their wings, or have bristles or other structures to link their wings together during flight.
The larvae are specialised predators, with elongated mandibles adapted for piercing and sucking. The larval body form varies between different families, depending on the nature of their prey. In general, however, they have three pairs of thoracic legs, each ending in two claws. The abdomen often has adhesive discs on the last two segments.
Life cycle and ecology
The larvae of most families are predators. Many chrysopids, hemerobids and coniopterygids eat aphids and other pest insects, and some have been used for biological control (either from commercial distributors, but also abundant and widespread in nature).
Larvae in various families cover themselves in debris (sometimes including dead prey insects) as camouflage, taken to an extreme in the ant lions, which bury themselves completely out of sight and ambush prey from "pits" in the soil. Larvae of some Ithonidae are root feeders, and larvae of Sisyridae are aquatic, and feed on freshwater sponges. A few mantispids are parasites of spider egg sacs.
As in other holometabolic orders, the pupal stage is enclosed in some form of cocoon composed of silk and soil or other debris. The pupa eventually cuts its way out of the cocoon with its mandibles, and may even move about for a short while before undergoing the moult to the adult form.
Adults of many groups are also predatory, but some do not feed, or consume only nectar.
The osmylids are of Jurassic or Early Cretaceous origin and may be the most ancient of the Neuropteran groups. The extinct osmylid Protosmylus is fossilized in middle Eocene Baltic amber. The genus Burmaleon is described from two fossils of Cenomanian age Burmese amber, implying crown group radiation in the Early Cretaceous or earlier. The family Kalligrammatidae lived from the Jurassic to Aptian (Lower Cretaceous) periods.
Molecular analysis in 2018 using mitochondrial rRNA and mitogenomic data places the Neuroptera within the Neuropterida, sister to the Raphidioptera and containing the Megaloptera (sister to the rest of the Neuroptera). The fossil record has contributed to the understanding of the group's phylogeny. Relationships within the Myrmeleontiformia are still in flux.
- Genus Mantispidiptera Grimaldi, 2000 (Late Cretaceous; New Jersey; formerly Mantispidae)
- Genus Mesohemerobius Ping, 1928(Late Jurassic/Early Cretaceous; China)
- Family Permithonidae † (probably paraphyletic)
- Family Prohemerobiidae † (probably paraphyletic)
- Family Nevrorthidae[Note 1]
- Family Grammosmylidae †
- Family Osmylitidae † (probably paraphyletic)
- Family Osmylidae: osmylids
- Superfamily Ithonioidea
- Family Ithonidae: moth lacewings (includes Rapismatidae and Polystoechotidae)
- Superfamily Chrysopoidea
- Superfamily Hemerobioidea
- Family Hemerobiidae: brown lacewings
- Superfamily Coniopterygoidea
- Superfamily Mantispoidea
- Superfamily Nemopteroidea
- Superfamily Myrmeleontoidea
- Family Osmylopsychopidae †
- Family Nymphitidae †
- Family Solenoptilidae † (probably paraphyletic)
- Family Brogniartiellidae †
- Family Nymphidae: split-footed lacewings (includes Myiodactylidae)
- Family Babinskaiidae †
- Family Myrmeleontidae: antlions (includes Palaeoleontidae)
- Family Ascalaphidae: owlflies, ascalaphids
In human culture
Five species of Neuroptera are among 1681 insect species eaten by humans worldwide.
- "Neurorthidae" is a lapsus.
- David Grimaldi & Michael S. Engel (2005). Evolution of the Insects. Cambridge University Press. ISBN 0-521-82149-5.
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- Michael S. Engel (2005). "A remarkable kalligrammatid lacewing from the Upper Jurassic of Kazakhstan (Neuroptera: Kalligrammatidae)". Transactions of the Kansas Academy of Science. 108 (1): 59–62. doi:10.1660/0022-8443(2005)108[0059:ARKLFT]2.0.CO;2.
- Breitkreuz, L. C. V.; Winterton, S. L.; Engel, M. S. (2017). "Wing tracheation in Chrysopidae and other Neuropterida (Insecta): a resolution of the confusion about vein fusion". American Museum Novitates. 3890: 1–44. doi:10.1206/3890.1.
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- Yan, Y.; Wang Y, Liu, X.; Winterton, S. L.; Yang, D. (2014). "The First Mitochondrial Genomes of Antlion (Neuroptera: Myrmeleontidae) and Split-footed Lacewing (Neuroptera: Nymphidae), with Phylogenetic Implications of Myrmeleontiformia". Int J Biol Sci. 10 (8): 895–908. doi:10.7150/ijbs.9454. PMC 4147223.CS1 maint: multiple names: authors list (link)
- Engel, Michael S.; Grimaldi, David A. (2007). "The neuropterid fauna of Dominican and Mexican amber (Neuropterida, Megaloptera, Neuroptera)" (PDF). American Museum Novitates (3587): 1–58.
- Myskowiak, J.; Huang, D.; Azar, D.; Cai, C.; Garrouste, R.; Nel, A. (2016). "New lacewings (Insecta, Neuroptera, Osmylidae, Nymphidae) from the Lower Cretaceous Burmese amber and Crato Formation in Brazil". Cretaceous Research. 59: 214. doi:10.1016/j.cretres.2015.10.029.
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- Yue, Bi-Song; Song, Nan; Lin, Aili; Zhao, Xincheng (2018). "Insight into higher-level phylogeny of Neuropterida: Evidence from secondary structures of mitochondrial rRNA genes and mitogenomic data". PLOS ONE. 13 (1): e0191826. doi:10.1371/journal.pone.0191826. ISSN 1932-6203.
- Grimaldi, D. A. & Engel, M. S., 2005: Evolution of the Insects. Cambridge University Press, 2005, pages xv-755
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- Parker, S. P. (ed.), 1982: Synopsis and classification of living organisms. Vols. 1 & 2. McGrew-Hill Book Company
- Jones, J.R. (2019) Total‐evidence phylogeny of the owlflies (Neuroptera, Ascalaphidae) supports a new higher‐level classification. Zoologica Scripta: 06 October 2019 https://doi.org/10.1111/zsc.12382
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- MacClancy, Jeremy (2007). Consuming the Inedible: Neglected Dimensions of Food Choice. Berghahn.
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- Illustrated database of Neuroptera (insects)
- A database of Neuroptera related scientific literature
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