Archaeobalbini Viidalepp, 1981

Murillo-Ramos, Leidys, Brehm, Gunnar, Sihvonen, Pasi, Hausmann, Axel, Holm, Sille, Ghanavi, Hamid Reza, Õunap, Erki, Truuverk, Andro, Staude, Hermann, Friedrich, Egbert, Tammaru, Toomas & Wahlberg, Niklas, 2019, A comprehensive molecular phylogeny of Geometridae (Lepidoptera) with a focus on enigmatic small subfamilies, PeerJ 7, pp. 1-39 : 24-25

Archaeobalbini Viidalepp, 1981 , status revised

(original spelling: Archeobalbini, justified emendation in Hausmann (1996))

Type genus: Archaeobalbis Prout, 1912 (synonymized with Herochroma Swinhoe, 1893 in Holloway (1996))

Material examined: Herochroma curvata Han & Xue, 2003 , H. baba Swinhoe 1893 , Metallolophia inanularia Han & Xue, 2004 , M. cuneataria Han & Xue, 2004 , Actenochroma muscicoloraria (Walker, 1862) , Absala dorcada Swinhoe, 1893 , Metaterpna batangensis Hang & Stüning, 2016 , M. thyatiraria (Oberthür, 1913) , Limbatochlamys rosthorni Rothschild, 1894 , Psilotagma pictaria (Moore, 1888) , Dindica para Swinhoe, 1893 , Dindicodes crocina (Butler, 1880) , Lophophelma erionoma (Swinhoe, 1893) , L. varicoloraria (Moore, 1868) , L. iterans (Prout, 1926) and Pachyodes amplificata (Walker, 1862) .

This lineage splits into four groups: Herochroma Swinhoe, 1893 + Absala Swinhoe, 1893 + Actenochroma Warren, 1893 is the sister lineage of the rest of Archaeobalbini that were recovered as three clades with unresolved relationships comprising the genera Limbatochlamys Rothschild, 1894 , Psilotagma Warren, 1894 , Metallolophia Warren, 1895 , Metaterpna Yazaki, 1992 , Dindica Warren, 1893 , Dindicodes Prout, 1912 , Lophophelma Prout, 1912 and Pachyodes Guenée, 1858 . This tribe can be diagnosed by the combination of DNA data from six genetic markers, see for instance Pachyodes amplificata CAD ( MG015522 ), COI ( MG014818 ), EF1a ( MG015409 ), GAPDH ( MG014941 ), MDH ( MG015057 ) and RpS5 ( MG015638 ). Branch support values in IQ-TREE confirm the monophyly of this clade (SH-like = 88.3, UFBoot2 = 64). GenBank accession numbers are shown in Supplementary Material. Amorphological diagnosis requires further research.

Xenozancla Warren, 1893 (unnamed G3) is sister to the clade comprising Dysphaniini and Pseudoterpnini s.str. Sihvonen et al. (2011) did not include Xenozancla in their analyses and suggested a sister relationship of Dysphaniini and Pseudoterpnini , but with low support. According to Ban et al. (2018), Xenozancla is more closely related to Pseudoterpnini s.str. than to Dysphaniini . However, due to low support, Ban et al. (2018) did not propose a taxonomic assignment for Xenozancla , which is currently not assigned to a tribe. Although our IQ-TREE results show that Xenozancla is sister to a clade comprising Dysphaniini and Pseudoterpnini s.str., the RAxML analysis did not recover the same phylogenetic relationships. Instead, Dysphaniini + Pseudoterpnini s.str. are found to be sister taxa, but Xenozancla is placed close to Rhomborista monosticta (Wehrli, 1924) . As in Ban et al. (2018), our results do not allow us to reach a conclusion about the phylogenetic affinities of these tribes, due to low support of nodes.

The Australian genus Crypsiphona Meyrick, 1888 (unnamed G4) was placed close to Hemitheini . Crypsiphona has been assigned to Pseudoterpnini (e. g. Pitkin, Han & James, 2007, Õunap & Viidalepp, 2009), but is recovered as a separate lineage in our tree. Given the isolated position of Crypsiphona , the designation of a new tribe could be considered, but due to low support of nodes in our analyses, further information (including morphology) is needed to confirm the phylogenetic position of this genus. In our phylogenetic hypothesis, a large clade including the former tribes Lophochoristini , Heliotheini , Microloxiini , Thalerini , Rhomboristini , Hemistolini , Comostolini , Jodini and Thalassodini is recovered as sister to the rest of Geometrinae . These results are in full agreement with Ban et al. (2018), who synonymized all of these tribes with Hemitheini .

Although the monophyly of Hemitheini is strongly supported, our findings recovered only a few monophyletic subtribes. For example, genera placed in Hemitheina were intermixed with those belonging to Microloxiina, Thalassodina and Jodina. Moreover, many genera which were unassigned to tribe, were recovered as belonging to Hemitheini . Our findings recovered Lophostola Prout, 1912 as sister to all Hemitheini . These results are quite different from those found by Ban et al. (2018) who suggested Rhomboristina as being sister to the rest of Hemitheini . In contrast, our results recovered Rhomboristina mingled with Hemistolina.

These different results are probably influenced by the presence of African and Madagascan Lophostola in our analysis. In our opinion the subtribe concept, as applied in Hemitheini earlier, is not practical and we do not advocate its use in geometrid classification.