Loss of male secondary sexual structures in allopatry in the Neotropical butterfly genus <I>Arcas</I> (Lycaenidae: Theclinae: Eumaeini)

Abstract

Male secondary sexual characters in Lepidoptera may be present or absent in species that otherwise appear to be closely related, an observation that has led to differences of opinion over the taxonomic usefulness of these structures above the species level. An evolutionary issue raised by this debate is whether male secondary sexual characters (1) can be regained after being lost evolutionarily, (2) are not lost after being evolved, or (3) are `switched on and off&#39; by genes that regulate development. A second evolutionary issue is the conditions under which male secondary sexual characters might be lost or gained evolutionarily. Because these structures are thought to promote species recognition, theory predicts evolutionary losses to be most likely in allopatry; evolutionary gains to be most likely during the process of secondarily establishing sympatry or during sympatric speciation. We updated the species-level taxonomy of the brilliant emerald-winged Neotropical lycaenid butterfly genus Arcas and performed an analysis of phylogenetic relations among species to assess these evolutionary issues. We morphologically detail a scent pouch on the ventral hindwing of Arcas and report that six species possess the pouch with androconia, one possesses the pouch without androconia, and the remaining two species have neither pouch nor androconia. In addition, eight Arcas species have a morphologically species-specific male forewing scent pad, and one lacks a scent pad. This variation appears to be the result of three evolutionary losses and no gains of male secondary sexual organs. The four Arcas species lacking a scent pouch or a scent pad are allopatric with their closest phylogenetic relatives while four of five with both of these structures are sympatric. Although Arcas is a small genus, these results are significantly more extreme than predicted by chance. For taxonomy, this study provides a rationale for the evolutionary loss of male secondary sexual structures and suggests that their absence, but itself, does not indicate a lack of relationship above the species level.