JODITI

JODITI + APORANDRIA + THALASSODITI

The grouping of the Joditi , Aporandria and Thalassoditi is well supported in this study. The subtribe Joditi (as Jodini) was established by Inoue (1961) and was treated as a provisional tribe, including four Palaearctic and Indo-Pacific genera ( Jodis , Berta , Gelasma and Thalerura Swinhoe , the latter two genera as synonyms of Maxates ) by Hausmann (1996). Viidalepp (1996) and Holloway (1996) included Joditi in the wider concept of Hemitheini . Han & Xue (2011a) followed this definition, which is also supported in this molecular study.

The species of Maxates number over 100 and were separated in Gelasma Warren , Maxates and Thalerura (Prout, 1912, 1920–41) until Holloway (1996) synonymized Gelasma and Thalerura with Maxates and summarized the most distinctive feature of Maxates as the possession of a strong hindwing tail and a ventral flap on the valva.

When Prout (1934 –38) described Ecchloropsis , he made a comparison with Hemistola and Dyschloropsis but did not mention Gelasma and Maxates , perhaps because he did not examine the male genitalia. Han & Xue (2011a) summarized the diagnostic characters of Ecchloropsis and Maxates and listed the differences between them. Although they noted that Ecchloropsis has a ventral flap in the valva similar to that found in typical Maxates , the authors did not synonymize the two, given that Ecchloropsis lacks a frenulum, whereas almost all species of Maxates have one.

In this analysis, the genus Maxates is found to be polyphyletic, as Maxates acutissima perplexata and M. grandificaria are sister to Ecchloropsis with full support, whereas another three species, M. sp.1, M. sp.2 and M. thetydaria , are grouped as sister to Berta and Jodis . The genus Berta is also found to be paraphyletic, since two species of Jodis are nested within Berta species (including the type species B. chrysolineata Walker ). Further research, including the type species of Maxates and Jodis and a greater number of other species, is needed to resolve the phylogenetic relationship among these four genera, the monophyly of the Joditi and the monophyly of Maxates and Berta .

The subtribe Thalassoditi (as Thalassodini) was also erected by Inoue (1961) with the inclusion of the genus Thalassodes Guenée , from which three genera ( Orothalassodes Holloway , Pelagodes Holloway and Remiformvalva Inoue ) were split and erected by Holloway (1996) and Inoue (2006), mainly based on the structures of the male genitalia and the eighth segment. Hausmann (1996) summarized the differential features of Thalassoditi as follows: cell of hindwing very short, discocellular vein oblique and lacking abdominal crests. Hausmann et al. (2016b) synonymized Thalassoditi with Hemistoliti because they share many morphological characters. However, the monophyly of Thalassoditi is fully supported in the present study, with Thalassodes at the basal position of the clade. The genera Prasinocyma and Albinospila Holloway are closely allied to Orothalassodes ( Holloway, 1996) , and they are expected to be clustered with Orothalassodes if they are sampled. Hausmann et al. (2016a) suggested that the current generic combinations within the large Thalassodes complex ( Holloway, 1996) need to be revised. Further research is needed to explore the internal structure of the Thalassoditi by including more genera, such as Prasinocyma and Albinospila .

Aporandria was placed in Hemistoliti by Hausmann (1996). He also stated that the external appearance of Aporandria resembles some genera of Geometrini but differs from the latter with respect to the tympanum and venation. In this study, the genus Aporandria is grouped with Thalassoditi with good support. The close relationship between Aporandria and Thalassoditi was first found in this study. However, we hesitate to place Aporandria in Thalassoditi due to its distinct wing patterns. Genera in Thalassoditi , together with Aporandria , are sister to part of Joditi and fall within Joditi . We suspect that the subtribe Thalassoditi may be a synonym of Joditi and that Aporandria also belongs to Joditi , but additional data are needed to validate this hypothesis.