Phylogenetic and Morphological Analysis of Wing Base Articulation in Vespidae (Hymenoptera): A Cladistic Approach

Insect wing base sclerites are crucial to wing function and evolution, yet their diversity beyond order-level comparisons remains poorly understood. We examine variation in wing base sclerites across Vespidae, focusing on the axillary sclerites (1Ax, 2Ax, and 3Ax), the shoulder sclerite, and associated structures. The first axillary sclerite shows distinct regional differentiation and bears a well-sclerotized knob that influences wing articulation. Additionally, 2Ax in Vespidae is a single, triangular structure with three attachment points, distinct from the two-part composition in some other wasps, which facilitates high-frequency wing vibrations. Our findings also highlight variable fusion patterns in 3Ax and its interaction with 2Ax, contributing to wing flexibility. The basiradial bridge, connecting the subcostal and radial veins, reinforces wing stability and articulation. Phylogenetic analysis based on wing-base morphology does not support the monophyly of Vespidae and differs from molecular hypotheses, but it refines previous morphological interpretations. The well-supported subfamily relationships confirm Vespinae as a monophyletic group and reveal a close association among Polistinae, Stenogastrinae, and Eumeninae, as represented by Polistes, Eustenogaster, and Oreumenes, respectively, suggesting evolutionary transitions in social behavior within the family Vespidae. The absence of a fourth axillary sclerite challenges earlier hypotheses, providing new insights into Hymenopteran wing base evolution. Two articulation models are proposed for forewings and hindwings, supported by three-dimensional reconstructions of axillary sclerites, indirect and direct flight muscles, and their attachment sites. These results refine interpretations of wasp wing mechanics, evolution, and morphological diversification across taxa.