Abstract

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.

Highlights

  • The genus Epicephala Meyrick, 1880 (Gracillariidae) has recently become an important taxon in ecology and evolutionary biology because of their mutualisms with plants in the genera Glochidion, Breynia and Phyllanthus (Phyllanthaceae) (Kato et al 2003; Kawakita and Kato 2009; Kawakita 2010)

  • Glochidion, Breynia and Phyllanthus belong to the well-defined tribe Phyllantheae, which contains over 1,200 species globally (Hoffmann et al 2006)

  • The purpose of this paper is to provide descriptions of the Epicephala species occurring in Japan, where most published ecological studies have been conducted

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Summary

Introduction

The genus Epicephala Meyrick, 1880 (Gracillariidae) has recently become an important taxon in ecology and evolutionary biology because of their mutualisms with plants in the genera Glochidion, Breynia and Phyllanthus (Phyllanthaceae) (Kato et al 2003; Kawakita and Kato 2009; Kawakita 2010). Epicephala moths are the only documented pollinators for many of these hosts, making this interaction an obligate mutualism. Because there is high level of species-specificity between the plants and the moths (Kawakita and Kato 2006; Kawakita et al 2010), a comparable number of Epicephala species likely exist; the genus currently consists of only 53 named species (De Prins and De Prins 2005, 2014; Li and Yang 2015; Li et al 2015). There is clearly a need to rapidly advance the taxonomy of the genus, especially because most published ecological and evolutionary studies on this mutualism treated Epicephala with arbitrary and variable names (Kawakita and Kato 2004a,b, 2009; Kawakita et al 2004, 2010, 2015; Okamoto et al 2007; Hembry et al 2012, 2013; Goto et al 2010; Svensson et al 2010), making comparisons among studies problematic (but see Hu et al 2011; Zang et al 2012a; Li and Yang 2015; Li et al 2015 for recent taxonomic advancement of Chinese Epicephala)

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