EVOLUTION OF THE HIND WING IN COLEOPTERA

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Abstract
A survey is made of the major features of the venation, articulation, and folding in the hind wings of Coleoptera. The documentation is based upon examination of 108 Coleoptera families and 200 specimens, and shown in 101 published figures. Wing veins and articular sclerites are homologized with elements of the neopteran wing groundplan, resulting in wing vein terminology that differs substantially from that generally used by coleopterists. We tabulate the differences between currently used venational nomenclature and the all-pterygote homologous symbols. The use of the neopteran groundplan, combined with the knowledge of the way in which veins evolved, provides many strong characters linked to the early evolutionary radiation of Coleoptera. The order originated with the development of the apical folding of the hind wings under the elytra executed by the radial and medial loop. The loops, which are very complex venational structures, further diversified in four distinctly different ways which mark the highest (suborder) taxa. The remaining venation and the wing articulation have changed with the loops, which formed additional synapomorphies and autapomorphies at the suborder, superfamily, and sometimes even family and tribe levels. Relationships among the four currently recognized suborders of Coleoptera are reexamined using hind wing characters. The number of wing-related apomorphies are 16 in Coleoptera, seven in Archostemata + Adephaga–Myxophaga, four in Adephaga–Myxophaga, seven in Myxophaga, nine in Archostemata, and five in Polyphaga. The following phylogenetic scheme is suggested: Polyphaga [Archostemata (Adephaga + Myxophaga)]. Venational evidence is given to define two major lineages (the hydrophiloid and the eucinetoid) within the suborder Polyphaga. The unique apical wing folding mechanism of beetles is described. Derived types of wing folding are discussed, based mainly on a survey of recent literature. A sister group relationship between Coleoptera and Strepsiptera is supported by hind wing evidence.