Some four-winged insect orders, such as the Lepidoptera, have developed a wide variety of morphological wing coupling mechanisms in the imago which render these taxa as "functionally dipterous" (effectively two-winged) for efficient insect flight. [1] All but the most basal forms exhibit this wing coupling. [2] : 4266
The mechanisms are of three different types - jugal, frenulo-retinacular and amplexiform. [3]
The more primitive groups of moth have an enlarged lobe-like area near the basal posterior margin, i.e. at the base of the forewing, called jugum, that folds under the hindwing in flight. [2] [4] : 631–664
Other groups of moth have a frenulum on the hindwing that hooks under a retinaculum on the forewing. [4] The retinaculum is a hook or tuft on the underside of the forewing of some moths. Along with the frenulum, a spine at the base of the forward or costal edge of the hindwing, it forms a coupling mechanism for the front and rear wings of the moth.
In the butterflies [a] and in the Bombycoidea, [b] there is no arrangement of frenulum and retinaculum to couple the wings. Instead, an enlarged humeral area of the hindwing is broadly overlapped by the forewing. Despite the absence of a specific mechanical connection, the wings overlap and operate in phase. The power stroke of the forewing pushes down the hindwing in unison. This type of coupling is a variation of frenate type but where the frenulum and retinaculum are completely lost. [1] [5]
Hamuli wing coupling is a mechanism unique to winged Hymenoptera. The hamuli consist of hook-like setae arrayed along the leading edge of the hind wing that engage with the recurved trailing edge (the retinaculum) of the fore wing during flight, uniting both wings into a single functional wing. [6]