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]
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