“…Therefore, the medulla has been the most plausible candidate where the first stage of color processing takes place (Kelber, 2016). In line with this assumption, spectral opponency, a key feature in color processing, had been reported in the medulla and/or higher visual centers in several insect species (Kelly & Mote, 1990;Kien & Menzel, 1977;Osorio, 1986;Paulk, Phillips-Portillo, Dacks, Fellous, & Gronenberg, 2008;Yang, Lin, & Hung, 2004). Rather unexpectedly, photoreceptors showing spectral opponency have also been found even at the level of the retina in butterflies (Chen, Arikawa, & Yang, 2013;Horridge, Mar celja, Jahnke, & Mati c, 1983;Mati c, 1983).…”