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Novel traits in the order Lepidoptera include prolegs in the abdomen of larvae, scales, and eyespot and band color patterns in the wings of adults. We review recent work that investigates the developmental origin and diversification of these four traits from a gene-regulatory network (GRN) perspective. While prolegs and eyespots appear to derive from distinct ancestral GRNs co-opted to novel body regions, scales derive from in situ modifications of a sensory bristle GRN. The origin of the basal and central symmetry systems of bands on the wing is associated with the expression of theWntAgene in those regions, whereas the more marginal bands depend on two other genes,Distal-lessandspalt. Finally, several genes have been discovered that play important roles in regulating background wing color, via the regulation of pigmentation GRNs. The identification of shared and novelcis-regulatory elements of genes belonging to these distinct GRNs helps trace the developmental and evolutionary history of these traits. Future work should examine the extent to which ancestral GRNs are co-opted/modified to produce the novel traits and how these GRNs map to specific cell types in ancestral and derived traits. 

The underlying genetic changes that regulate the appearance and disappearance of repeated traits, or serial homologs, remain poorly understood. One hypothesis is that variation in genomic regions flanking master regulatory genes, also known as input–output genes, controls variation in trait number, making the locus of evolution almost predictable. Another hypothesis implicates genetic variation in up- or downstream loci of master control genes. Here, we use the butterfly Bicyclus anynana , a species that exhibits natural variation in eyespot number on the dorsal hindwing, to test these two hypotheses. We first estimated the heritability of dorsal hindwing eyespot number by breeding multiple butterfly families differing in eyespot number and regressing eyespot numbers of offspring on midparent values. We then estimated the number and identity of independent genetic loci contributing to eyespot number variation by performing a genome-wide association study with restriction site-associated DNA sequencing from multiple individuals varying in number of eyespots sampled across a freely breeding laboratory population. We found that dorsal hindwing eyespot number has a moderately high heritability of ∼0.50 and is characterized by a polygenic architecture. Previously identified genomic regions involved in eyespot development, and novel ones, display high association with dorsal hindwing eyespot number, suggesting that homolog number variation is likely determined by regulatory changes at multiple loci that build the trait, and not by variation at single master regulators or input–output genes.