NEMORIINI

Ban, Xiaoshuang, Jiang, Nan, Cheng, Rui, Xue, Dayong & Han, Hongxiang, 2018, Tribal classification and phylogeny of Geometrinae (Lepidoptera: Geometridae) inferred from seven gene regions, Zoological Journal of the Linnean Society 184 (3), pp. 653-672 : 667-668

NEMORIINI + SYNCHLORINI + COMIBAENINI

Nemoriini (as Nemoriinae in Gumppenberg, 1887) is a New World tribe including many genera, with Ochrognesiini as a representative in the IndoAustralian tropics. The tribe Ochrognesiini was established by Inoue (1961) to include the two genera Chloromachia Warren and Ochrognesia Warren , both of which were synonymized with Eucyclodes Warren , together with six other genera by Holloway (1996). Accordingly, Ochrognesiini was synonymized with Nemoriini . The diagnostic features of Nemoriini were summarized and commented on by Pitkin (1996), Hausmann (1996) and Holloway (1996). Most recently, Viidalepp (2017) recognized 25 genera in Nemoriini and summarized three main features: the rod-shaped uncus of the male genitalia, the specific shape of the eighth abdominal sternite of the male abdomen and larvae with unclothed chalazae.

Ferguson (1969) first erected Synchlorini and placed the genera Synchlora Guenée , Merochlora Prout (= Synchlora ), Cheteoscelis Prout (= Synchlora ) in it. Synchlorini is characterized by male genitalia in which the uncus is reduced and the socii are rigidly sclerotized, tapered and pointed ( Pitkin, 1996).

The tribe Comibaenini is a small-sized group in the Geometrinae and was first established by Inoue (1961), including the genera Comibaena Hübner ( Chlorochromodes Warren and Comostolodes Warren as synonyms, and Colutoceras Warren as a subgenus) and Thetidia Boisduval. Hausmann (1996) added two new genera, Microbaena Hausmann and Proteuchloris Hausmann , and suggested that the tribe is monophyletic. Holloway (1996) recognized Comibaenini (as subtribe Comibaeniti ) as a discrete group. He included the genera Comibaena , Argyrocosma Turner , Comostolodes (with Chlorochromodes and Hercoloxia as synonyms of Comostolodes ) and Thetidia , and added a new genus, Protuliocnemis Holloway. Han, Galsworthy & Xue (2012) reviewed Comibaenini worldwide, embracing eight genera: Comibaena , Microbaena , Thetidia , Proteuchloris , Linguisaccus Han et al. , Chlorochromodes , Argyrocosma and Protuliocnemis . The most distinctive feature of Comibaenini is the bifid vinculum of the male genitalia. This study sampled five known genera of Comibaenini and the monophyly of Comibaenini is well supported. Within the tribe, Protuliocnemis + Argyrographa are sister to Linguisaccus + ( Comibaena + Thetidia ). Comibaena and Thetidia are closely related, and Comibaena is paraphyletic, in that Th. chlorophyllaria (Hedemann) and Th. albocostaria (Bremer) are nested within Comibaena , indicating that these two species may be transferred to Comibaena . Han et al. (2012) discussed the similarity among some species of Comibaena ( C. hypolampes Prout , C. cenocraspis Prout , C. latilinea Prout and C. swanni Prout ) and Thetidia , stating that the frenulum is present in Comibaena species but absent in Thetidia . Prout (1932: 20) regarded the absence or presence of a frenulum in the male as an important character at the level of genus and above. However, Pitkin (1996) stated that both states can occur within a genus, giving as examples Synchlora , Oospila , Chloractis Warren and Phrudocentra Warren. Therefore , it is possible to include Thetidia in Comibaena . Given the morphological diversity of Comibaena members, it is also possible that it needs to be split into different genera.

Previous studies have shown a close relationship among Nemoriini , Synchlorini and Comibaenini . Ferguson (1985) stated that Synchlorini have much more in common with Nemoriini , and he would have included the Synchlorini in Nemoriini , were it not for differences in genitalia, venation and larval behaviour. In contrast, Pitkin (1993) stated that the genital differences between Synchlorini and Nemoriini appear insufficient to justify separating tribal status, and deferred formal synonymy due to a lack of information on the early stages of some other important genera, such as Lissochlora Warren and Chavarriella Pitkin. Later, Pitkin (1996) mentioned that some genera in Nemoriini share a similar valva structure with Synchlora (Synchlorini) and Chlorissa (Hemitheini) , and continued using Synchlorini , adding Xenopepla Warren to it. Holloway (1996) stated that Synchlorini are probably related to Comibaenini , based on the larval habit of attaching debris and some aspects of genitalic structure. In summary, these three tribes are closely related, as shown by their similar larval behaviour and some genitalic features.

In this analysis, Nemoriini , Synchlorini and Comibaenini form a monophyletic clade with strong support. The tribe Synchlorini is clustered with Nemoria Hübner with full support, and Comibaenini is recovered as sister to Pyrochlora with moderate support. A similar result was found by Sihvonen et al. (2011), unsurprisingly, as the data for Nemoriini ( Nemoria , Pyrochlora ) and Synchlorini ( Synchlora ) used in the current study were mainly obtained from their research. The monophyly of Nemoriini is not supported by the present concept, under which it would be a monophyletic group with the inclusion of Synchlorini and Comibaenini . Considering that only five species in three genera of Nemoriini are included in our analysis, and there are more than 130 species in Nemoria alone, we defer a decision on this synonymy at present. Further analysis, including additional taxa, is needed to determine the relationship among these tribes and the position of Pyrochlora .