title: Forewing color pattern in Micropterigidae (Insecta: Lepidoptera): homologies between contrast boundaries, and a revised hypothesis for the origin of symmetry systems.
abstract: Despite the great importance of lepidopteran wing patterns in various biological disciplines, homologies between wing pattern elements in different moth and butterfly lineages are still not understood. Among other reasons, this may be due to an incomplete understanding of the relationship between color pattern and wing venation; many individual wing pattern elements have a known relationship with venation, but a framework to unite all wing pattern elements with venation is lacking. Though plesiomorphic wing veins are known to influence color patterning even when not expressed in the adult wing, most studies of wing pattern evolution have focused on derived taxa with a reduced suite of wing veins.
  
  
  
  The present study aims to address this gap through an examination of Micropterigidae, a very early-diverged moth family in which all known plesiomorphic lepidopteran veins are expressed in the adult wing. The relationship between wing pattern and venation was examined in 66 species belonging to 9 genera. The relationship between venation and pattern element location, predicted based on moths in the family Tortricidae, holds for Sabatinca just as it does for Micropterix. However, the pattern elements that are lightly colored in Micropterix are dark in Sabatinca, and vice-versa. When plotted onto a hypothetical nymphalid wing in accordance with the relationship between pattern and venation discussed here, the wing pattern of Sabatinca doroxena very closely resembles the nymphalid groundplan.
  
  
  
  The color difference in pattern elements between Micropterix and Sabatinca indicates that homologies exist among the contrast boundaries that divide wing pattern elements, and that color itself is not a reliable indicator of homology. The similarity between the wing pattern of Sabatinca doroxena and the nymphalid groundplan suggests that the nymphalid groundplan may have originated from a Sabatinca-like wing pattern subjected to changes in wing shape and reduced expression of venation.
authors.affiliation: Mississippi Entomological Museum, Mississippi State, MS, 39762, USA. schachatsr@si.edu.|Department of Paleobiology, Smithsonian Institution, Washington, DC, 20013, USA. schachatsr@si.edu.
authors.forename: Sandra R
authors.initials: SR
authors.lastname: Schachat
authors.affiliation: Mississippi Entomological Museum, Mississippi State, MS, 39762, USA.
authors.forename: Richard L
authors.initials: RL
authors.lastname: Brown
mesh_terms: D000818:Animals; D005075:Biological Evolution; D003116:Color; D009036:Moths / Q000033:anatomy & histology*; D014680:Veins / Q000033:anatomy & histology; D014921:Wings, Animal / Q000033:anatomy & histology*
keywords: Developmental constraints; Microlepidoptera; Nymphalid groundplan; Symmetry systems