Abstract

The nymphalid groundplan (NGP) has proven to be extraordinarily useful in the study of butterfly color patterns because it allows for the identification of homologous elements across species. It has long been thought that the NGP is broadly applicable to all Lepidoptera, implying that the characters which make-up butterfly color patterns are homologous to those found in the moths. However, this conjecture remains mostly untested. We analyzed the wing patterns of the hyper-diverse arctiid tiger moths, which are represented worldwide by approximately 11,000 species, and found that the color patterns of these animals can be parsed into a limited toolkit of homologous characters. Some of the pattern elements identified, such as the basal and central symmetry system, are present on the NGP, but their morphology is often quite different from what is seen in the butterflies. The border ocelli of the NGP appear to be absent altogether in the Arctiidae, and conversely, two distal pattern element we term the ‘van Bemmelen’ and ‘terminal’ bands are present in the color patterns of many arctiids, but are not represented on the NGP. In light of the observed differences, we derive a new theoretical groundplan based on the original NGP that we refer to as the ‘arctiid archetype.’ This model provides a comprehensive framework for understanding how the wing patterns of these animals develop, and yields novel insight into their evolutionary history.

Highlights

  • The wing patterns of butterflies have proven to be a uniquely tractable system for research at the interface of evolution and development (Nijhout, 1991; Beldade and Brakefield, 2002; Jiggins, 2017)

  • The aim of this paper is to broaden our understanding of lepidopteran wing patterning by determining whether the dorsal forewing patterns of the hyperdiverse arctiid “tiger moths” [Lepidoptera, Erebidae (Arctiidae)]1 can be decomposed into an underlying system of characters that are homologous to elements found on the nymphalid groundplan

  • We suggest that the dorsal forewing patterns of the arctiids are constructed from a maximum of five primary characters: a basal and central symmetry system, along with a discal spot, and two external characters we have termed the “van Bemmelen,” and “terminal” bands

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Summary

Introduction

The wing patterns of butterflies have proven to be a uniquely tractable system for research at the interface of evolution and development (Nijhout, 1991; Beldade and Brakefield, 2002; Jiggins, 2017). A theoretical archetype known as the nymphalid groundplan (NGP) (Nijhout, 1991) allows for the identification of these homologous characters within and across species, making it possible to precisely describe the results of experiments designed to interfere with pattern development (Monteiro et al, 2006; Martin and Reed, 2014; Mazo-Vargas et al, 2017), and identify large-scale trends in pattern evolution (Monteiro, 2008). A previous study has shown that it is possible to locate NGP characters such as the basal symmetry system, central symmetry system, and discal spot in the patterns of the tiger moth Utetheisa ornatrix (Gawne and Nijhout, 2019), but no attempt has yet been made to assess the generality of this finding through a large-scale study of the Arctiidae

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