Adult neuropterans have four membranous wings, all about the same size, with manyveins. They have chewing mouthparts, and undergo completemetamorphosis.
Neuropterans first appeared during thePermianperiod, and continued to diversify through theMesozoic era.[2] During this time, several unusually large forms evolved, especially in the extinctfamilyKalligrammatidae, often called "thebutterflies of theJurassic" for their large, patterned wings.[3]
Neuropterans are soft-bodied insects with relatively few specialised features. They have large lateralcompound eyes, and may or may not also haveocelli. Their mouthparts have strongmandibles suitable for chewing, and lack the various adaptations found in most otherholometabolan insect groups.
They have four wings, usually similar in size and shape, and a generalised pattern of veins.[4] Some neuropterans have specialised sense organs in their wings, or have bristles or other structures to link their wings together during flight.[5]
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. Theabdomen often has adhesive discs on the last two segments.[5] There is no connection between the midgut and the hindgut in the larvae, and their Malpighian tubules are instead used for producing silk for their cocoon, which exits through their anus. In the families Myrmeleontidae and Ascalaphidae the tenth abdominal segment is modified into a spinneret that is usually retracted into the ninth abdominal segment, and used for the construction of the cocoon.[6]
Larvae in various families cover themselves in debris (including other insects, living and dead[9]) ascamouflage, taken to an extreme in theant lions, which bury themselves completely out of sight and ambush prey from "pits" in the soil. Larvae of someIthonidae are root feeders, and larvae ofSisyridae are aquatic, and feed onfreshwatersponges. A fewmantispids areparasites ofspider egg sacs.
As in otherholometabolic orders, thepupal stage is enclosed in some form ofcocoon 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 themoult to the adult form.[5]
Adults of many groups are also predatory, but some do not feed, or consume onlynectar.
Molecular analysis in 2018 using mitochondrial rRNA and mitogenomic data places the Megaloptera assister to Neuroptera, and Raphidioptera as sister to this combined lineage, though these results were considered tentative.[17][10] Thefossil record has contributed to the understanding of the group's phylogeny.[1][18][19][20] Relationships within theMyrmeleontiformia are still in flux.[21]
A phylogenomic analysis published in 2023 confirmed the topology of the neuropterid orders and found the relationships between the families of Neuropterida as shown in the following phylogenetic tree.[22]
Review of the Neropterid group orders by Engel, Winterton, and Breitkreuz (2018) included grouping of the Neuropteran families in a nested set of clades, an abandonment of the paraphyletic suborder "Hemerobiiformia" and redefinition of Myrmeleontiformia as a clade.[23]
The fossil genus †MesohemerobiusPing, 1928 from the Late Jurassic–Early Cretaceous of China has been treated asincertae sedis within Neuroptera, while the fossil families †Permoberothidae and †Permithonidae are treated as a sister group to clade Eidoneuroptera formed by Neuroptera + Megaloptera.[23]
The use of Neuroptera inbiological control of insect pests has been investigated, showing that it is difficult to establish and maintain populations in fields of crops.[25]
Five species of Neuroptera are among 1681 insect species eaten by humans worldwide.[26]
The New Guinea Highland people claim to be able to maintain a muscular build and great stamina despite their low protein intake as a result ofeating insects including Neuroptera.[27]
^abcHoell, H. V., Doyen, J. T. & Purcell, A. H. (1998).Introduction to Insect Biology and Diversity, 2nd ed. Oxford University Press. pp. 447–450.ISBN0-19-510033-6.{{cite book}}: CS1 maint: multiple names: authors list (link)
^Powell, Erin. (2023). Defensive behaviors of the mealybug Nipaecoccus nipae (Maskell, 1893) (Hemiptera: Pseudococcidae) and the green lacewing Ceraeochrysa claveri (Navás, 1911) (Neuroptera: Chrysopidae), with videos of dorsal packet loading and mealybug ostiole function. Insecta Mundi. 1–11.
^Bechly, G.; Makarkin, V. N. (2016). "A new gigantic lacewing species (Insecta: Neuroptera) from the Lower Cretaceous of Brazil confirms the occurrence of Kalligrammatidae in the Americas".Cretaceous Research.58:135–140.Bibcode:2016CrRes..58..135B.doi:10.1016/j.cretres.2015.10.014.
^Grimaldi, D. A. & Engel, M. S., 2005: Evolution of the Insects. Cambridge University Press, 2005, pages xv-755
^Engel, M. S. & Grimaldi, D. A., 2007: The neuropterid fauna of Dominican and Mexican amber (Neuropterida: Megaloptera, Neuroptera).American Museum Novitates: #3587, pages 1-58
^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 2019https://doi.org/10.1111/zsc.12382
^abEngel, M. S.; Winterton, S. L.; Breitkreuz, L. C. (2018). "Phylogeny and evolution of Neuropterida: where have wings of lace taken us?".Annual Review of Entomology.63:531–551.doi:10.1146/annurev-ento-020117-043127.PMID29324039.
^Engel, M. S.; Grimaldi, D. A. (2008). "Diverse Neuropterida in Cretaceous amber, with particular reference to the paleofauna of Myanmar (Insecta)".Nova Supplementa Entomologica.20:1–86.
^Xu, X. X. (2014). "Electrophysiological and Behavior Responses of Chrysopa phyllochroma (Neuroptera Chrysopidae) to Plant Volatiles".Environmental Entomology.44 (5):1425–1433.doi:10.1093/ee/nvv106.ISSN0046-225X.PMID26314008.S2CID46558266.
^Ramos-Elorduy, J. (2005). Maurizio G. Paoletti (ed.).Insects: a hopeful resource. Enfield, New Hampshire: Science Publishers. pp. 263–291.ISBN978-1-57808-339-8.{{cite book}}:|work= ignored (help)
^MacClancy, Jeremy (2007).Consuming the Inedible: Neglected Dimensions of Food Choice. Berghahn.