Revision of the Japanese species of Epicephala Meyrick with descriptions of seven new species (Lepidoptera, Gracillariidae)

Epicephala moths are involved in obligate mutualisms with their Phyllanthaceae hosts, in which the female moths assure pollination and, in return, their progeny develop by consuming the seeds. Ecological, molecular and geographical data suggest that the genus includes several hundred species, but the majority remains to be formally described. Here we revise the Japanese species of Epicephala Meyrick, 1880. In addition to two previously named species, seven species are newly described: Epicephala anthophilia sp. n., Epicephala lanceolatella sp. n., Epicephala perplexa sp. n., Epicephala obovatella sp. n., Epicephala corruptrix sp. n., Epicephala parasitica sp. n. and Epicephala nudilingua sp. n. The first four are species involved in obligate pollination mutualism, while the fifth is a pollinating seed parasite and the last two are derived non-pollinating seed parasites of herbaceous Phyllanthus. Each of the nine Japanese Epicephela species is specialized to a single plant species in the genera Glochidion, Breynia or Phyllanthus, except for Epicephala obovatella and Epicephala corruptrix that each utilizes two closely related Glochidion species. Considerable variations are found in pollination and oviposition behaviors among species, which are reflected in their proboscis and ovipositor morphologies, respectively. Molecular phylogeny indicated that there have been repeated transitions in oviposition mode during the diversification of Epicephala, which were accompanied by changes in ovipositor morphology, as suggested by a correlation analysis. Keys to species are provided.