Wing pattern diversity in Brassolini butterflies (Nymphalidae, Satyrinae)

Abstract

This study describes and compares the diverse dorsal and ventral wing color patterns of Brassolini butterflies. Thirty-three species are illustrated, where pattern elements of the nymphalid ground plan are labeled in color. In general, a larger number of pattern elements can be identified on the ventral than on the dorsal surface of both wings, and the forewing has a larger number of discernible pattern elements than the hind wing. The dorsal elements are broad, diffuse, and more difficult to identify against the typically brown brassoline wing background color. Species with a light colored dorsal background served as a guide for our proposal that fewer pattern elements are present dorsally, particularly on the hind wing. Colorful bands or markings generally present on the dorsal surface seem to be associated with specific pattern elements and have correspondence to the ventral pattern. We refer to these as trailing bands, and they constitute a predominant feature of the brassoline dorsal coloration. We propose a subordinate groundplan for brassolines and interpret some of the ventral pattern variation in light of their phylogeny. Dorsal color pattern variation that leads to sexual dimorphism and mimetic resemblance are also discussed. Resumo: Este estudo descreve e compara os diversos padrões de coloração dorsal e ventral das borboletas da tribo Brassolini. Com o objetivo de demarcar os elementos do padrão de base dos ninfalídeos ('pattern elements of the nymphalid ground plan'), 33 espécies são ilustradas. De maneira geral, o número de elementos-padrão identificados na face ventral é maior do que o da face dorsal em ambas as asas, e a asa anterior contém um número maior de elementos visíveis do que a posterior. Os elementos da face dorsal são mais largos e difusos, e também mais difíceis de identificar devido à usual coloração de fundo marrom escuro dos brassolíneos. Espécies que apresentam coloração de fundo clara serviram como guia para a nossa interpretação de que a superfície dorsal das asas apresenta um número reduzido de elementos-padrão, especialmente a da asa posterior. Faixas ou manchas coloridas que geralmente aparecem na superfície dorsal estão aparentemente associadas com elementos-padrão específicos, e apresentam correspondência com faixas que aparecem na face ventral. Estas faixas ('trailing bands') se alastram a partir da borda de certos elementos e constituem uma característica predominante da coloração dorsal dos brassolíneos. Nós propomos um padrão de base para os brassolíneos que é subordinado ao dos ninfalídeos, e utilizamos a filogenia do grupo para interpretar alguns aspectos da variação de elementos-padrão da superfície ventral. As variações de padrão e cor que resultam em dimorfismo sexual e mimetismo são discutidas. Palavras-chave: borboleta, coloração das asas, padrão de base dos ninfalídeos, manchas ocelares, ocelo, Caligo. This study represents the first broad survey of color pattern diversity in Brassolini. Here we provide a framework for understanding color pattern varia...

Symmetry systems on the wings ofDichromodesGuenée (Lepidoptera: Geometridae) are unconstrained by venation

“…Of note, the predictions of these two hypotheses overlap somewhat: each predicts that the distal edge of the central symmetry system reaches the costa proximal to the distalmost branch of the Subcostal vein (Figs. 1, 2B and 2D), a prediction that dates to the inception of the NGP (Schwanwitsch, 1924;Süffert, 1927) and is supported by recent studies (Nijhout, 1991;Otaki, 2012;Penz & Mohammadi, 2013).…”

“…Full-size DOI: 10.7717/peerj.8263/ fig-1 butterfly genera such as Bicyclus Kirby, 1871 (Robertson & Monteiro, 2005), Junonia Hübner [1819] (Kodandaramaiah, 2009), Heliconius Kluk, 1802 (Reed & Gilbert, 2004), Papilio Linnaeus, 1758 (Kunte et al, 2014), Pieris Schrank 1801 (Takayama & Yoshida, 1997), and Vanessa Fabricius 1807 (Abbasi & Marcus, 2015) are disproportionately represented among the Lepidoptera that have been successfully reared in a laboratory setting. Furthermore, traditional morphological (Otaki, 2012;Penz & Mohammadi, 2013;Taira et al, 2015) and physiological (Kusaba & Otaki, 2009;Nijhout, 2010) approaches also typically focus on butterflies alone. The popularity of moths in the family Saturniidae has increased in recent years for studies that focus on wing pattern along the termen or within the interior of the wing (Marcus, 2019;Monteiro et al, 2006;Sourakov, 2017).…”

Documenting diversity in the Amazonian butterfly genus Bia (Lepidoptera, Nymphalidae)

“…(1) MF DFW white apical ocelli: These are part of the border ocellus series (Nijhout 1991, Penz & Mohammadi 2013, and their size varies continuously within and between taxa. In combination with other characters, this character was diagnostic for one species.…”

“…(7) F VFW ripple pattern of the postmedial area: The ripple pattern can be similar to the remainder of the VFW or broadly spaced, revealing a pale yellow color. Pattern element f of the nymphalid groundplan (Nijhout 1991;Penz & Mohammadi 2013) can be visible or inconspicuous. This varies locally, and also geographically within taxa.…”

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

“…2.1, 2.2 and 2.7). In many Brassolini and also Morpho, the hind wing ocelli are clearly positioned in the medial area of the wing, which can produce a striking visual effect depending on their size (Penz and Mohammadi 2013;Figs. 2.1d-e and 2.2c).…”