Insect wing

Original veins and wing posture of a dragonfly
Hoverflies hovering to mate
A cockchafer's hardened forewings raised, hindwings unfolding
Outstretched wing of a bumblebee

Insect wings are adult outgrowths of the insect exoskeleton that enable insects to fly. They are found on the second and third thoracic segments (the mesothorax and metathorax), and the two pairs are often referred to as the forewings and hindwings, respectively, though a few insects lack hindwings, even rudiments. The wings are strengthened by a number of longitudinal veins, which often have cross-connections that form closed "cells" in the membrane (extreme examples include the dragonflies and lacewings). The patterns resulting from the fusion and cross-connection of the wing veins are often diagnostic for different evolutionary lineages and can be used for identification to the family or even genus level in many orders of insects.

Physically, some insects move their flight muscles directly, others indirectly. In insects with direct flight, the wing muscles directly attach to the wing base, so that a small downward movement of the wing base lifts the wing itself upward. Those insects with indirect flight have muscles that attach to and deform the thorax, causing the wings to move as well.

The wings are present in only one sex (often the male) in some groups such as velvet ants and Strepsiptera, or are selectively lost in "workers" of social insects such as ants and termites. Rarely, the female is winged but the male not, as in fig wasps. In some cases, wings are produced only at particular times in the life cycle, such as in the dispersal phase of aphids. Wing structure and colouration often vary with morphs, such as in the aphids, migratory phases of locusts and polymorphic butterflies. At rest, the wings may be held flat, or folded a number of times along specific patterns; most typically, it is the hindwings which are folded, but in a few groups such as the vespid wasps, it is the forewings.

The evolutionary origin of the insect wing is debated. During the 19th century, the question of insect wing evolution originally rested on two main positions. One position postulated insect wings evolved from pre-existing structures, while the second proposed insect wings were entirely novel formations.[1][2] The “novel” hypothesis suggested that insect wings did not form from pre-existing ancestral appendages but rather as outgrowths from the insect body wall.[3]

Long since, research on insect wing origins has built on the “pre-existing structures” position that was originally proposed in the 19th century.[2] Recent literature has pointed to several ancestral structures as being important to the origin of insect wings. Among these include: gills, respiratory appendages of legs, and lateral (paranotal) and posterolateral projections of the thorax to name a few.[4]

According to more current literature, possible candidates include gill-like structures, the paranotal lobe, and the crustacean tergal plate. The latter is based on recent insect genetic research which indicates that insects are pan-crustacean arthropods with a direct crustacean ancestor and shared genetic mechanisms of limb development.[3][5][6][7][8]

Other theories of the origin of insect wings are the paranotal lobe theory, the gill theory and the dual theory of insect wing evolution. These theories postulate that wings either developed from paranotal lobes, extensions of the thoracic terga;[5] that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies;[5] or that insect wings arose from the fusion of pre-existing endite and exite structures each with pre-existing articulation and tracheation.[9][10]

  1. ^ Crampton, G. (1916). "The Phylogenetic Origin and the Nature of the Wings of Insects According to the Paranotal Theory". Journal of the New York Entomological Society. 24 (1): 1–39. JSTOR 25003692.
  2. ^ a b Ross, Andrew (2017). "Insect Evolution: The Origin of Wings". Current Biology. 27 (3): R113–R115. doi:10.1016/j.cub.2016.12.014. PMID 28171756 – via Web of Science.
  3. ^ a b Averof, Michalis, and S. M. Cohen. (1997). "Evolutionary origin of insect wings from ancestral gills". Nature. 385 (6617): 627–630. Bibcode:1997Natur.385..627A. doi:10.1038/385627a0. PMID 9024659. S2CID 4257270 – via Web of Science.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  4. ^ Grodnitsky, Dmitry, L. (1999). Form and Function of Insect Wings: The Evolution of Biological Structures. pp. 82–83.{{cite book}}: CS1 maint: multiple names: authors list (link)
  5. ^ a b c Alexander, David, E. (2015). On the Wing: Insects, Pterosaurs, Birds, Bats and the Evolution of Animal Flight. Oxford University Press. pp. 74–101.{{cite book}}: CS1 maint: multiple names: authors list (link)
  6. ^ Haug, Joachim, C. Haug., and R. J. Garwood. (2016). "Evolution of insect wings and development – new details from Palaeozoic nymphs". Biological Reviews. 91 (1): 53–69. doi:10.1111/brv.12159. PMID 25400084. S2CID 21031689.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  7. ^ Almudi, Isabel; Vizueta, Joel; Wyatt, Christopher D. R.; de Mendoza, Alex; Marlétaz, Ferdinand; Firbas, Panos N.; Feuda, Roberto; Masiero, Giulio; Medina, Patricia; Alcaina-Caro, Ana; Cruz, Fernando (2020). "Genomic adaptations to aquatic and aerial life in mayflies and the origin of insect wings". Nature Communications. 11 (1): 2631. Bibcode:2020NatCo..11.2631A. doi:10.1038/s41467-020-16284-8. ISSN 2041-1723. PMC 7250882. PMID 32457347.
  8. ^ Bruce, Heather, N.H. Patel. (2020). "Knockout of crustacean leg patterning genes suggests that insect wings and body walls evolved from ancient leg segments". Nature Ecology & Evolution. 4 (12): 1703–1712. Bibcode:2020NatEE...4.1703B. doi:10.1038/s41559-020-01349-0. PMID 33262517. S2CID 227253368.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  9. ^ Clark-Hatchel, Courtney (2013). "Insights into insect wing origin provided by functional analysis of vestigial in the red flour beetle, Tribolium castaneum". Proceedings of the National Academy of Sciences of the United States of America. 110 (42): 16951–16956. Bibcode:2013PNAS..11016951C. doi:10.1073/pnas.1304332110. PMC 3801059. PMID 24085843.
  10. ^ Prokop, Jakub, Pecharová, M., Nel, A., Hörnschemeyer, T., Krzemińska, E., Krzemiński, W., & Engel, M (2017). "Paleozoic Nymphal Wing Pads Support Dual Model of Insect Wing Origins". Current Biology. 27 (2): 263–269. doi:10.1016/j.cub.2016.11.021. PMID 28089512.{{cite journal}}: CS1 maint: multiple names: authors list (link)

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