The nymphalid groundplan is an archetypical color pattern of nymphalid butterflies involving three major symmetry systems and a discal symmetry system, which share the basic morphogenesis unit. Here, the morphological and spatial relationships among these symmetry systems were studied based on cross-species comparisons of nymphalid hindwings. Based on findings in Neope and Symbrenthia, all three major symmetry systems can be expressed as bands, spots, or eyespot-like structures, suggesting equivalence (homology) of these systems in developmental potential. The discal symmetry system can also be expressed as various structures. The discal symmetry system is circularly surrounded by the central symmetry system, which may then be surrounded by the border and basal symmetry systems, based mainly on findings in Agrias, indicating a unified supersymmetry system covering the entire wing. The border symmetry system can occupy the central part of the wing when the central symmetry system is compromised, as seen in Callicore. These results suggest that butterfly color patterns are hierarchically constructed in a self-similar fashion, as the fractal geometry of the nymphalid groundplan. This self-similarity is likely mediated by the serial induction of organizers, and symmetry breaking of the system morphology may be generated by the collision of opposing signals during development.
Keywords: Lepidoptera; Nymphalidae; butterfly wing; color pattern; color pattern element; fractal; nymphalid groundplan; organizer; self-similarity; serial induction; symmetry breaking; symmetry system.