Exploring Color Pattern Diversification in Early Lineages of Satyrinae (Nymphalidae)

Abstract

Based on the most recent nymphalid phylogeny, the Satyrinae can be tentatively organized into the species-rich tribe Satyrini plus a clade that includes the Morphini, Brassolini, Haeterini, Elymniini, Melanitini, Dirini, Zetherini, and Amathusiini. Members of the latter eight tribes have the largest body sizes within Satyrinae and also show extraordinary wing pattern variation. Representatives of these tribes are illustrated herein, and pattern elements of the nymphalid ground plan are identified. Five themes are briefly discussed in light of their pattern diversification: (1) central symmetry system dislocations, (2) variation in ventral hind wing ocelli, (3) the color band between elements f and g, (4) sexual dimorphism and mimicry, and (5) transparency. Within an ecological and evolutionary standpoint, selected genera are provided as examples to explore wing patterns involved in male mating displays, camouflage, and mimicry.

Morphological and Spatial Diversity of the Discal Spot on the Hindwings of Nymphalid Butterflies: Revision of the Nymphalid Groundplan

“…Pierellization occurs because each wing compartment is an independent unit of color pattern development [ 24 , 25 ]. Identifications of elements and importance of pierellization can be seen in some studies [ 26 , 27 , 28 , 29 ]. Moreover, in the forewings, the discal spot and the central symmetry system are probably often more complicated with surrounding elements than in the hindwings.…”

“…As in previous studies [ 3 , 4 , 5 , 6 , 7 , 8 , 26 , 27 , 28 , 29 ], identification processes for elements were based on a “homology search” between different symmetry systems in the same wing surface, between closely related species, and between different individuals of the same species. Identification of the discal spot was basically made in reference to wing veins and to the distal and proximal bands of the central symmetry system (dBC and pBC), provided that these bands were expressed.…”

The Fractal Geometry of the Nymphalid Groundplan: Self-Similar Configuration of Color Pattern Symmetry Systems in Butterfly Wings

“…Fifth, recent examples of homology searches for elemental identification in butterfly wings have also been found elsewhere [ 9 , 12 , 33 , 34 , 35 , 36 , 37 ]. Among these cases, the genus Cethosia , in which white bands are present not only between elements but also between subelements within a symmetry system, is particularly interesting [ 9 ].…”

“…Finally, the assumption of homologous relationship among the symmetry systems has been widely accepted in many studies of color patterns as noted above [ 2 , 3 , 4 , 5 , 8 , 9 , 10 , 11 , 12 , 23 , 24 , 25 , 27 , 28 , 33 , 34 , 35 , 36 , 37 , 38 , 39 ], which is also supported by the concept of the secondary symmetry systems [ 2 ]. However, I acknowledge that it is difficult to completely exclude an alternative possibility that morphological similarities among the symmetry systems evolved via convergence from very different evolutionary origins.…”