Like many other members of the insect world, the lift generated by butterflies is more than what can be accounted for by steady-state, non-transitory aerodynamics. Studies using Vanessa atalanta in a windtunnel show that they use a wide variety of aerodynamic mechanisms to generate force. These include wake capture, vortices at the wing edge, rotational mechanisms and Weis-Fogh 'clap-and-fling' mechanisms. The butterflies were also able to change from one mode to another rapidly. Insects are the only group of invertebrates known to have evolved flight. Insects possess some remarkable flight characteristics and abilities, still far superior to attempts by humans to replicate their capabilities. Even our understanding of the aerodynamics of flexible, flapping wings and how insects fly is imperfect. One application of this research is in the engineering of extremely small micro air vehicles with low Reynolds numbers. 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. Insect wings do not constitute appendages in technical parlance, as insects only have one pair of appendages per segment. 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 Odonata and Neuroptera). 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 identifica ion to the family or even genus level in many orders of insects. The physical dynamics of flight are composed of direct and indirect flight. Those species that employ direct flight have wing muscles directly attached the wing base so that a small movement of the wing base downward, lifts the wing itself upward. However Insects with indirect flight have muscles that attach to thorax and deform it; since the wings are extensions of the thoracic exoskeleton, the deformations of the thorax cause the wings to move as well. The wings may be present in only one sex (often the male) in some groups such as velvet ants and Strepsiptera, or 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. Beyond the mere presence/absence of wings, the structure and colouration will often vary with morphs, such as in the aphids, migratory phases of locusts and in 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 very few groups such as vespid wasps, it is the forewings. How and why insect wings evolved is not well understood. Three main theories on the origins of insect flight are that wings developed from paranotal lobes, extensions of the thoracic terga; that they are modifications of movable abdominal gills as found on aquatic naiads of mayflies and that insect wings arose from the fusion of pre-existing endite and exite structures each with pre-existing articulation and tracheation.