identifier	taxonID	type	CVterm	format	language	title	description	additionalInformationURL	UsageTerms	rights	Owner	contributor	creator	bibliographicCitation
038F87AD2B0EFFA52EC55929FB2908B2.text	038F87AD2B0EFFA52EC55929FB2908B2.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Larentiinae Duponchel 1845	<div><p>Larentiinae Duponchel, 1845</p> <p>Larentiinae are a monophyletic entity (Fig. 3). In concordance with the results of Sihvonen et al. (2011), Viidalepp (2011), Õunap, Viidalepp &amp; Truuverk (2016) and Strutzenberger et al. (2017), Dyspteridini are supported as sister to all other larentiines. Remarkably, Brabirodes Warren, 1904 forms an independent lineage. Chesiadini are monophyletic and sister to all larentiines except Dyspteridini, Brabirodes and Trichopterygini. These results do not support the suggestion by Viidalepp (2006) and Sihvonen et al. (2011) that Chesiadini are sister to Trichopterygini.</p> <p>In our phylogenetic hypothesis, Asthenini are sister to the Perizomini + Melanthiini + Eupitheciini clade. These results do not fully agree with Õunap, Viidalepp &amp; Truuverk (2016) who found Asthenini to be sister to all Larentiinae except Dyspteridini, Chesiadini, Trichopterygini and Eudulini. However, our results do support the Melanthiini + Eupitheciini complex as a sister lineage to Perizomini. Sihvonen et al. (2011) recovered Phileremini and Rheumapterini as well-supported sister taxa. Our results suggest Triphosa dubitata Linnaeus 1758 (Triphosini) is sister to Phileremini, with Rheumapterini sister to this clade. Cidariini were recovered as paraphyletic, as the genera Coenotephria Prout, 1914 and Lampropteryx Stephens, 1831 cluster in a different clade (unnamed clade L7) apart from the lineage comprising the type genus of the tribe, Cidaria Treitschke, 1825.</p> <p>Ceratodalia Packard, 1876, currently placed in Hydriomenini and Trichodezia Warren, 1895 are nested within Cidariini. These results are not in concordance with Õunap, Viidalepp &amp; Truuverk (2016), who regarded this tribe to be monophyletic. Scotopterygini are sister to a lineage comprising Ptychorrhoe blosyrata Guenée (1858), Disclisioprocta natalata (Walker, 1862) (placed in the unnamed clade L8), Euphyiini, an unnamed clade L9 comprising the genera Pterocypha, Archirhoe and Obila, Xanthorhoini and Cataclysmini. Euphyiini are monophyletic, but Xanthorhoini are recovered as mixed with Cataclysmini. The same findings were shown by Õunap, Viidalepp &amp; Truuverk (2016), but no taxonomic rearrangements were proposed. Larentiini are monophyletic and sister of Hydriomenini, Heterusiini, Erateinini, Stamnodini and some unnamed clades (L11–14). Although with some differences, our results support the major phylogenetic patterns of Õunap, Viidalepp &amp; Truuverk (2016).</p> <p>Despite substantial progress, the tribal classification and phylogenetic relationships of Larentiinae are far from being resolved (Õunap, Viidalepp &amp; Truuverk, 2016). Forbes (1948) proposed eight tribes based on morphological information, Viidalepp (2011) raised the number to 23 and Õunap, Viidalepp &amp; Truuverk (2016) recovered 25 tribes studying 58 genera. Our study includes 23 of the currently recognized tribes and 125 genera (with an emphasis on Neotropical taxa). However, the phylogenetic position of many taxa remains unclear, and some tropical genera have not yet been formally assigned to any tribe. Formal descriptions of these groups will be treated in detail by G. Brehm et al. (2019, unpublished data) and E. Õunap et al. (2019, unpublished data).</p> </div>	http://treatment.plazi.org/id/038F87AD2B0EFFA52EC55929FB2908B2	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B01FFA92EC55D59FC7A0919.text	038F87AD2B01FFA92EC55D59FC7A0919.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Orthostixinae Meyrick 1892	<div><p>Orthostixinae Meyrick, 1892</p> <p>Orthostixinae were not included in our study. Sihvonen et al. (2011) showed this subfamily as deeply embedded within Ennominae, but unfortunately it was not represented by the type genus of the subfamily. These results agree with Holloway (1996) who examined Orthostixis Hübner, (1823) and suggested the inclusion in Ennominae despite the full development of hindwing vein M2, the presence of a forewing areole and the very broad base of the tympanal ansa. We sampled the species Naxa textilis (Walker, 1856) and Orthostixis cribraria (Hübner, 1799), but only three and one marker were successfully sequenced for these samples, respectively. We included these species in the preliminary analyses but results were so unstable that we excluded them from the final analysis. Further research including fresh material and more genetic markers are needed to investigate the position of Orthostixinae conclusively.</p> </div>	http://treatment.plazi.org/id/038F87AD2B01FFA92EC55D59FC7A0919	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B0AFFA22EC55E9BF9C40B6F.text	038F87AD2B0AFFA22EC55E9BF9C40B6F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Epidesmiinae Murillo-Ramos, Brehm & Sihvonen 2019	<div><p>Epidesmiinae Murillo-Ramos, Brehm &amp; Sihvonen newsubfamily</p> <p>LSIDurn:lsid:zoobank.org:act:34D1E8F7-99F1-4914-8E12-0110459C2040</p> <p>Type genus: Epidesmia Duncan &amp; Westwood, 1841.</p> <p>Material examined: Taxa included in the molecular phylogeny: Ecphyas holopsara Turner, 1929, Systatica xanthastis Lower, 1894, Adeixis griseata Hudson, 1903, Dichromodes indicataria Walker, 1866, Phrixocomes sp. Turner, 1930, Abraxaphantes perampla Swinhoe, 1890, Epidesmia chilonaria (Herrich-Schäffer, 1855), Phrataria replicataria Walker, 1866.</p> <p>Most of the slender-bodied Oenochrominae, excluded from Oenochrominae s.str. by Holloway (1996), were recovered as an independent lineage (Fig. 4) that consists of two clades: Ec. holopsara + S. xanthastis and Ep. chilonaria + five other genera. Branch support values from IQ-TREE strongly support the monophyly of this clade (SH-like and UFBoot2 = 100), while in RAxML the clade is moderately supported (RBS = 89).</p> <p>These genera have earlier been assigned to Oenochrominae s.l. (Scoble &amp; Edwards, 1990). However, we recovered the group as a well-supported lineage independent from Oenochrominae s.str. and transfer them to Epidesmiinae, subfam. n. (Table 2).</p> <p>Phylogenetic position: Epidesmiinae is sister to Oenochrominae s.str. + Eumelea + Geometrinae + Ennominae.</p> <p>Short description of Epidesmiinae: Antennae in males unipectinate (exception: Adeixis), shorter towards the apex. Pectination moderate or long. Thorax and abdomen slender (unlike in Oenochrominae). Forewings with sinuous postmedial line and areole present. Forewings planiform (with wings lying flat on the substrate) in resting position, held like a triangle and cover the hindwings.</p> <p>Diagnosis of Epidesmiinae: The genera included in this subfamily form a strongly supported clade with DNA sequence data from the following gene regions (exemplar Epidesmia chilonaria (Herrich-Schäffer, 1855)) ArgK (MK738299), Ca-ATPase (MK738690), CAD (MK738960), COI (MK739187), EF1a (MK740168), GAPDH (MK740402), MDH (MK740974) and Nex9 (MK741433). Athorough morphological investigation of the subfamily, including diagnostic characters, is under preparation.</p> <p>Distribution: Most genera are distributed in the Australian region, with some species ranging into the Oriental region. Abraxaphantes occurs exclusively in the Oriental region.</p> </div>	http://treatment.plazi.org/id/038F87AD2B0AFFA22EC55E9BF9C40B6F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B0AFFA12EC55980FAE00E09.text	038F87AD2B0AFFA12EC55980FAE00E09.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Geometrinae Leach 1815	<div><p>Geometrinae Stephens, 1829</p> <p>The monophyly of Geometrinae is strongly supported, but the number of tribes included in this subfamily is still unclear. Sihvonen et al. (2011) analyzed 27 species assigned to 11 tribes, followed by Ban et al. (2018) with 116 species in 12 tribes. Ban et al. (2018) synonymized nine tribes, and validated the monophyly of 12 tribes, with two new tribes Ornithospilini and Agathiini being the first two clades branching off the main lineage of Geometrinae. Our study (168 species) validates the monophyly of 13 tribes, eleven of which were defined in previous studies: Hemitheini, Dysphaniini, Pseudoterpnini s.str., Ornithospilini, Agathiini, Aracimini, Neohipparchini, Timandromorphini, Geometrini, Comibaeini, Nemoriini. One synonymization is proposed: Synchlorini Ferguson, 1969 syn. nov. is synonymized with Nemoriini Gumppenberg, 1887. One tribe is proposed as new: Chlorodontoperini trib. nov., and one tribe (Archaeobalbini Viidalepp, 1981, stat. rev.) is raised from synonymy with Pseudoterpnini.</p> <p>Ban et al. (2018) found that Ornithospila Warren, 1894 is sister to the rest of Geometrinae, and Agathia Guenée, 1858 is sister to the rest of Geometrinae minus Ornithospila. Although weakly supported, our results (with more species of Agathia sampled) placed Ornisthospilini+Agathiini together and these tribes are the sister to the rest of Geometrinae. Chlorodontopera is placed as an isolated lineage as shown by Ban et al. (2018). Given that Chlorodontopera clearly forms an independent and well-supported lineage we propose the description of a new tribe Chlorodontoperini.</p> </div>	http://treatment.plazi.org/id/038F87AD2B0AFFA12EC55980FAE00E09	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B09FFA02EC55C29FCD50EFA.text	038F87AD2B09FFA02EC55C29FCD50EFA.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Chlorodontoperini , Murillo-Ramos, Sihvonen & Brehm 2019	<div><p>Chlorodontoperini Murillo-Ramos, Sihvonen &amp; Brehm, new tribe</p> <p>LSIDurn:lsid:zoobank.org:act:0833860E-A092-43D6-B2A1-FB57D9F7988D</p> <p>Type genus: Chlorodontopera Warren, 1893</p> <p>Material examined: Taxa in the molecular phylogeny: Chlorodontopera discospilata (Moore, 1867) and Chlorodontopera mandarinata (Leech, 1889).</p> <p>Some studies (Inoue, 1961; Holloway, 1996) suggested the morphological similarities of Chlorodontopera Warren, 1893 with members of Aracimini. Moreover, Holloway (1996) considered this genus as part of Aracimini. Our results suggest a sister relationship of Chlorodontopera with a large clade comprising Aracimini, Neohipparchini, Timandromorphini, Geometrini, Nemoriini and Comibaenini. Considering that our analysis strongly supports Chlorodontopera as an independent lineage (branch support SH-like = 99 UFBoot2 = 100, RBS = 99), we introduce the monobasic tribe Chlorodontoperini. This tribe can be diagnosed by the combination of DNA data from six genetic markers (exemplar Chlorodontopera discospilata) CAD (MG015448), COI (MG014735), EF1a (MG015329), GAPDH (MG014862), MDH (MG014980) and RpS5 (MG015562). Ban et al. (2018) did not introduce a new tribe because the relationship between Chlorodontopera and Euxena Warren, 1896 was not clear in their study. This relationship was also been proposed by Holloway (1996) based on similar wing patterns. Further analyses are needed to clarify the affinities between Chlorodontopera and Euxena.</p> <p>The tribe Chlorodontoperini is diagnosed by distinct discal spots with pale margins on the wings, which are larger on the hindwing; a dull reddish-brown patch is present between the discal spot and the costa on the hindwing, and veins M3 and CuA1 are not stalked on the hindwing (Ban et al., 2018). In the male genitalia, the socii are stout and setose and the lateral arms of the gnathos are developed, not joined. Sternite 3 of the male has setal patches (see Holloway, 1996 for illustrations). Formal taxonomic changesare listedin Table 2.</p> <p>Aracimini, Neohipparchini, Timandromorphini, Geometrini and Comibaenini were recovered as monophyletic groups. These results are in full agreement with Ban et al. (2018). However, the phylogenetic position of Eucyclodes Warren, 1894 is uncertain (unnamed G2). The monophyly of Nemoriini and Synchlorini is not supported. Instead, Synchlorini are nested within Nemoriini (support branch SH-like = 98.3, UFBoot2 = 91, RBS = 93). Our findings are in concordance with Sihvonen et al. (2011) and Ban et al. (2018), but our analyses included a larger number of markers and a much higher number of taxa. Thus, we formally synonymize Synchlorini syn. nov. with Nemoriini (Table 2).</p> <p>The monophyly of Pseudoterpnini sensu Pitkin, Han &amp; James (2007) could not be recovered. Similar results were shown by Ban et al. (2018) who recovered Pseudoterpnini s.l. including all the genera previously studied by Pitkin, Han &amp; James (2007), forming a separate clade from Pseudoterpna Hübner, 1823 + Pingasa Moore, 1887. Our results showed African Mictoschema Prout, 1922 falling within Pseudoterpnini s.str., and it is sister to Pseudoterpna and Pingasa. Asecond group of Pseudoterpnini s.l. was recovered as an independent lineage clearly separate from Pseudoterpnini s.str. (SH-like = 88.3, UFBoot2 = 64). Ban et al. (2018) did not introduce a new tribe due to the morphological similarities and difficulty in finding apomorphies of Pseudoterpnini s.str. In addition, their results were weakly supported. Considering that two independent studies have demonstrated the paraphyly of Pseudoterpnini sensu Pitkin et al. (2007), we see no reason for retaining the wide concept of this tribe. Instead, we propose the revival of the tribe status of Archaeobalbini.</p> </div>	http://treatment.plazi.org/id/038F87AD2B09FFA02EC55C29FCD50EFA	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B08FFAF2EC55B05FA4F0B3F.text	038F87AD2B08FFAF2EC55B05FA4F0B3F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Archaeobalbini Viidalepp 1981	<div><p>Archaeobalbini Viidalepp, 1981, status revised</p> <p>(original spelling: Archeobalbini, justified emendation in Hausmann (1996))</p> <p>Type genus: Archaeobalbis Prout, 1912 (synonymized with Herochroma Swinhoe, 1893 in Holloway (1996))</p> <p>Material examined: Herochroma curvata Han &amp; Xue, 2003, H. baba Swinhoe 1893, Metallolophia inanularia Han &amp; Xue, 2004, M. cuneataria Han &amp; Xue, 2004, Actenochroma muscicoloraria (Walker, 1862), Absala dorcada Swinhoe, 1893, Metaterpna batangensis Hang &amp; 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).</p> <p>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.</p> <p>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.</p> <p>The Australian genus Crypsiphona Meyrick, 1888 (unnamed G4) was placed close to Hemitheini. Crypsiphona has been assigned to Pseudoterpnini (e. g. Pitkin, Han &amp; James, 2007, Õunap &amp; 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.</p> <p>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.</p> <p>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.</p> </div>	http://treatment.plazi.org/id/038F87AD2B08FFAF2EC55B05FA4F0B3F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B07FFAB2EC55947FB400E30.text	038F87AD2B07FFAB2EC55947FB400E30.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Ennominae Duponchel 1845	<div><p>Ennominae Duponchel, 1845</p> <p>Ennominae are the most species-rich subfamily of geometrids. The loss of vein M2 on the hindwing is probably the best apomorphy (Holloway, 1994), although vein M2 is present as tubular in a few ennomine taxa (Staude, 2001; Skou &amp; Sihvonen, 2015). Ennominae are a morphologically highly diverse subfamily, and attempts to find further synapomorphies shared by all major tribal groups have failed.</p> <p>The number of tribes as well as phylogenetic relationships among tribes are still debated (see Skou &amp; Sihvonen, 2015 for an overview). Moreover, the taxonomic knowledge of this subfamily in tropical regions is still poor. Holloway (1994) recognized 21 tribes, Beljaev (2006) 24 tribes, and Forum Herbulot (2007) 27 tribes. To date, four molecular studies have corroborated the monophyly of Ennominae (Yamamoto &amp; Sota, 2007; Wahlberg et al., 2010; Õunap et al., 2011, Sihvonen et al., 2011), with Young (2006) being the only exception who found Ennominae paraphyletic. Moreover, four large-scale taxonomic revisions (without a phylogenetic hypothesis) were published by Pitkin (2002) for the Neotropical region, Skou &amp; Sihvonen (2015), Müller et al. (2019) for the Western Palaearctic region, and Holloway (1994) for Borneo. More detailed descriptions of taxonomic changes in Ennominae will be given by G. Brehm et al. (2019, unpublished data) and L. MurilloRamos et al. (2019, unpublished data). We here discuss general patterns and give details for taxonomic acts not covered in the other two papers.</p> <p>Our findings recover Ennominae as a monophyletic entity, but results were not highly supported in RAxML (RBS = 67) compared to IQ-TREE (SH-Like =100, UFBoot2 = 99). The lineage comprising Geometrinae and Oenochrominae is recovered as the sister clade of Ennominae. In previous studies, Wahlberg et al. (2010) sampled 49 species of Ennominae, Õunap et al. (2011) sampled 33 species, and Sihvonen et al. (2011) 70 species including up to eight markers per species. All these studies supported the division of Ennominae into “boarmiine” and “ennomine” moths (Holloway, 1994). This grouping was proposed by Forbes (1948) and Holloway (1994), who suggested close relationships between the tribes Boarmiini, Macariini, Cassymini and Eutoeini based on the bifid pupal cremaster and the possession of a fovea in the male forewing. The remaining tribes were defined as “ennomines” based on the loss of a setal comb on male sternum A3 and the presence of a strong furca in male genitalia. Both Wahlberg et al. (2010) and Sihvonen et al. (2011) found these two informal groupings to be reciprocally monophyletic.</p> <p>In our analyses, 653 species with up to 11 markers were sampled, with an emphasis on Neotropical taxa, which so far had been poorly represented in the molecular phylogenetic analyses. Our results recovered the division into two major subclades (Fig. 6), a core set of ennomines in a well-supported clade, and a poorly supported larger clade that includes the “boarmiines” among four other lineages usually thought of as "ennomines". The traditional “ennomines” are thus not found to be monophyletic in our analyses, questioning the utility of such an informal name. Our phylogenetic hypothesis supports the validation of numerous tribes proposed previously, in addition to several unnamed clades. We validate 23 tribes (Forum Herbulot, 2007; Skou &amp; Sihvonen, 2015): Gonodontini, Gnophini, Odontoperini, Nacophorini, Ennomini, Campaeini, Alsophilini, Wilemaniini, Prosopolophini, Diptychini, Theriini, Plutodini, Palyadini, Hypochrosini, Apeirini, Epionini, Caberini, Macariini, Cassymini, Abraxini, Eutoeini and Boarmiini.</p> <p>We hereby propose one new tribe: Drepanogynini trib. nov. (Table 2). Except for the new tribe, most of the groups recovered in this study are in concordance with previous morphological classifications (Holloway, 1994; Beljaev, 2006, 2016; Forum Herbulot, 2007; Skou &amp; Sihvonen, 2015; Müller et al., 2019).</p> <p>Five known tribes and two further unnamed lineages (E1, E2 in Fig. 6) form the core Ennominae: Gonodontini, Gnophini, Odontoperini, Nacophorini and Ennomini.</p> <p>Several Neotropical clades that conflict with the current tribal classification of Ennominae will be described as new tribes by G. Brehm et al. (2019, unpublished data). Gonodontini and Gnophini are recovered as sister taxa. Gonodontini was defined by Forbes (1948) and studied by Holloway (1994), who showed synapomorphies shared by Gonodontis Hübner, (1823), Xylinophylla Warren, 1898 and Xenimpia Warren, 1895. Our results recovered the genus Xylinophylla as sister of Xenimpia and Psilocladia Warren, 1898. Psilocladia is an African genus currently unassigned to tribe (see Sihvonen, Staude &amp; Mutanen, 2015 for details). Considering the strong support and that the facies and morphology are somewhat similar to other analyzed taxa in Gonodontini, we formally include Psilocladia in Gonodontini (Table 2). Gnophini are monophyletic and we formally transfer the African genera Oedicentra Warren, 1902 and Hypotephrina Janse, 1932, from unassigned to Gnophini (Table 2). The total number of species, and number of included genera in Gnophini are still uncertain (Skou &amp; Sihvonen, 2015; Müller et al., 2019). Based on morphological examination, Beljaev (2016) treated Angeronini as a synonym of Gnophini. The costal projection on male valva bearing a spine or group of spines was considered as a synapomorphy of the group. Using molecular data, Yamamoto &amp; Sota (2007) showed a close phylogenetic relationship between Angerona Duponchel, 1829 (Angeronini) and Chariaspilates Wehrli, 1953 (Gnophini). Similar results were shown by Sihvonen et al. (2011) who recovered Angerona and Charissa Curtis, 1826 as sister taxa, and our results also strongly support treating Angeronini as synonym of Gnophini.</p> <p>Holloway (1994) suggested close affinities among Nacophorini, Azelinini and Odontoperini on the basis of larval characters. In a morphology-based phylogenetic study, Skou &amp; Sihvonen (2015) suggested multiple setae on the proleg on A6 of the larvae as a synapomorphy of the group. Our results also support a close relationship of Nacophorini, Azelinini and Odontoperini. These clades will be treated in more detail by G. Brehmetal. (2019, unpublisheddata).</p> <p>Following the ideas of Pitkin (2002), Beljaev (2008) synonymized the tribes Ourapterygini and Nephodiini with Ennomini. He considered the divided vinculum in male genitalia and the attachment of muscles m 3 as apomorphies of the Ennomini, but did not provide a phylogenetic analysis. Sihvonen et al. (2011) supported Beljaev’ s assumptions and recovered Ennomos Treitschke, 1825 (Ennomini), Ourapteryx Leach, 1814 (Ourapterygini) and Nephodia Hübner, 1823 (Nephodiini) as belonging to the same clade. Our comprehensive analysis confirms those previous findings and we agree with Ennomini as the valid tribal name for this large clade. This clade will be treated in more detail by G. Brehm et al.</p> <p>(2019, unpublisheddata).</p> <p>Campaeini, Alsophilini, Wilemaniini and Prosopolophini grouped together in a well-supported clade (SH-like = 100, UFBoot2 = 99). Previous molecular analyses have shown an association of Colotoini [= Prosopolophini] and Wilemaniini (Yamamoto &amp; Sota, 2007; Sihvonen et al., 2011), although no synapomorphies are known to support synonymization (Skou &amp; Sihvonen, 2015). The Palaearctic genera Compsoptera Blanchard, 1845, Apochima Agassiz, 1847, Dasycorsa Prout, 1915, Chondrosoma Anker, 1854 and Dorsispina Nupponen &amp; Sihvonen, 2013, are potentially part of the same complex (Skou &amp; Sihvonen, 2015, Sihvonen pers. obs.), but they were not included in the current study.</p> <p>Campaeini is a small group including four genera with Oriental, Palaearctic and Nearctic distribution, apparently closely related to Alsophilini and Prosopolophini, but currently accepted as a tribe (Forum Herbulot, 2007; Skou &amp; Sihvonen, 2015). Our results support the close phylogenetic affinities among these tribes, but due to the limited number of sampled taxa, we do not propose any formal changes.</p> <p>The genus Declana Walker, 1858 is recovered as an isolated clade sister to Diptychini.</p> <p>This genus is endemic to New Zealand, but to date has not been assigned to tribe. According to our results, Declana could well be defined as its own tribe. However, the delimitation of this tribe is beyond the scope of our paper and more genera from Australia and New Zealand should first be examined. Aclose relationship between Nacophorini and Lithinini was suggested by Pitkin (2002), based on the similar pair of processes of the anellus in the male genitalia. Pitkin also noted a morphological similarity in the male genitalia (processes of the juxta) shared by Nacophorini and Diptychini. In a study of the Australasian fauna, Young (2008) suggested the synonymization of Nacophorini and Lithinini. This was further corroborated by Sihvonen, Staude &amp; Mutanen (2015) who found that Diptychini were nested within some Nacophorini and Lithinini. However, none of the studies proposed formal taxonomic changes because of limited taxon sampling. In contrast, samples in our analyses cover all biogeographic regions and the results suggest that true Nacophorini is a clade which comprises almost exclusively New World species. This clade is clearly separate from Old World “nacophorines” (cf. Young, 2003) that are intermixed with Lithinini and Diptychini. We here formally transfer Old World nacophorines to Diptychini and synonymize Lithinini syn. nov. with Diptychini (Table 2). Further formal taxonomic changes in the Nacophorini complex are provided by G. Brehm et al. (2019, unpublished data).</p> <p>Theria Hübner 1825, the only representative of Theriini in this study, clustered together with Lomographa Hübner, 1825 (Baptini in Skou &amp; Sihvonen, 2015), in a well-supported clade, agreeing with the molecular results of Sihvonen et al. (2011). The placement of Lomographa in Caberini (Rindge, 1979; Pitkin, 2002) is not supported by our study nor by that of Sihvonen et al. (2011). The monophyly of Lomographa has not been tested before, but we show that one Neotropical and one Palaearctic Lomographa species indeed group together. Our results show that Caberini are not closely related to the Theriini + Baptini clade, unlike in earlier morphology-based hypotheses (Rindge, 1979; Pitkin, 2002). Morphologically, Theriini and Baptini are dissimilar, therefore we recognize them as valid tribes (see description and illustrations in Skou &amp; Sihvonen, 2015).</p> <p>According to our results, 11 molecular markers were not enough to infer phylogenetic affinities of Plutodini (represented by one species of Plutodes). Similar results were found by Sihvonen et al. (2011), who in some analyses recovered Plutodes as sister of Eumelea. Our analyses are congruent with those findings. IQ-TREE results suggest that Plutodes is sister to Palyadini, but RAxML analyses recovered Eumelea as the most probable sister of Plutodes. Given that our analyses are not in agreement on the sister-group affinities of Plutodes, we do not make any assumptions about its phylogenetic position. Instead, we emphasize that further work needs to be done to clarify the phylogenetic positions of Plutodes and related groups.</p> <p>Hypochrosini is only recovered in a well-defined lineage if the genera Apeira Gistl, 1848 (Apeirini), Epione Duponchel, 1829 (Epionini), Sericosema (Caberini), Ithysia (Theriini), Capasa Walker, 1866 (unassigned) and Omizodes Warren, 1894 (unassigned) were transferred to Hypochrosini. Skou &amp; Sihvonen (2015) already suggested aclose association of Epionini, Apeirini and Hypochrosini. We think that synonymizing these tribes is desirable. However, due to the limited number of sampled taxa we do not propose any formal changes until more data becomes available. We do suggest, however, formal taxonomic changes for the genera Capasa and Omizodes from unassigned to Hypochrosini (Table 2).</p> <p>The southern African genus Drepanogynis is paraphyletic and has earlier been classified as belonging in Ennomini, and later in Nacophorini (Krüger, 2002). In our phylogeny, it is intermixed with the genera Sphingomima Warren, 1899, and Thenopa Walker, 1855.</p> <p>Hebdomophruda errans Prout, 1917 also clusters together with these taxa, apart from other Hebdomophruda Warren, 1897 species, which suggests that this genus is polyphyletic.</p> <p>These genera form a clade sister to the lineage that comprises several Hypochrosini species. Considering that our analysis strongly supports this clade, we place Thenopa, Sphingomima and Drepanogynis in a tribe of their own.</p> </div>	http://treatment.plazi.org/id/038F87AD2B07FFAB2EC55947FB400E30	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B03FFA92EC55C40FCFE0F18.text	038F87AD2B03FFA92EC55C40FCFE0F18.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Drepanogynini , Murillo-Ramos, Sihvonen & Brehm 2019	<div><p>Drepanogynini Murillo-Ramos, Sihvonen &amp; Brehm new tribe LSIDurn:lsid:zoobank.org:act:AA384988-009F-4175-B98C-6209C8868B93</p> <p>Type genus: Drepanogynis Guenée, (1858)</p> <p>The African genera Thenopa, Sphingomima and Drepanogynis appear as a strongly supported lineage (SH-like, UFBoot2 and RBS = 100). Krüger (1997, p. 259) proposed " Boarmiini and related tribes as the most likely sister group" for Drepanogynis, whereas more recently Drepanogynis was classified in the putative southern hemisphere Nacophorini (Krüger, 2014; Sihvonen, Staude &amp; Mutanen, 2015). In the current phylogeny, Drepanogynis is isolated from Nacophorini sensu stricto and from other southern African genera that have earlier been considered to be closely related to it (Krüger, 2014 and references therein). The other southern African genera appeared to belong to Diptychini in our study. The systematic position of Drepanogynis tripartita (Warren, 1898) has earlier been analyzed in a molecular study (Sihvonen, Staude &amp; Mutanen, 2015). The taxon grouped together with the Palaearctic species of the tribes Apeirini, Theriini, Epionini and putative Hypochrosini. Sihvonen, Staude &amp; Mutanen (2015) noted that Argyrophora trofonia (Cramer, 1779) (representing Drepanogynis group III sensu Krüger, 1999) and Drepanogynis tripartita (representing Drepanogynis group IV sensu Krüger, 2002) did not group together, but no formal changes were proposed. Considering that the current analysis strongly supports the placement of Drepanogynis and related genera in an independent lineage, and the aforementioned taxa in the sister lineage (Apeirini, Theriini, Epionini and putative Hypochrosini) have been validated at tribe-level, we place Drepanogynis and related genera in a tribe of their own.</p> <p>Material examined and taxa included: Drepanogynis mixtaria (Guenée, 1858), D. tripartita, D. determinata (Walker, 1860), D. arcuifera Prout, 1934, D. arcuatilinea Krüger, 2002, D. cnephaeogramma (Prout, 1938), D. villaria (Felder &amp; Rogenhofer, 1875), “ Sphingomima ” discolucida Herbulot, 1995 (genus combination uncertain, see taxonomic notes below), Thenopa diversa Walker, 1855, “ Hebdomophruda ” errans Prout, 1917 (genus combination uncertain, see taxonomic notes below).</p> <p>Taxonomic notes: We choose Drepanogynis Guenée, 1858 as the type genus for Drepanogynini, although it is not the oldest valid name (ICZN Article 64), because extensive literature has been published on Drepanogynis (Krüger, 1997, 1998, 1999, 2014), but virtually nothing exists on Thenopa, Walker, 1855, except the original descriptions of its constituent species. Current results show the urgent need for more extensive phylogenetic studies within Drepanogynini. Thenopa and Sphingomima are embedded within Drepanogynis, rendering it paraphyletic, but our taxon coverage is too limited to propose formal changes in this species-rich group. Drepanogynini, as defined here, are distributed in sub-Saharan Africa. Drepanogynis sensu Krüger (1997, 1998, 1999, 2014) includes over 150 species and it ranges from southern Africa to Ethiopia (Krüger, 2002, Vári, Kroon &amp; Krüger, 2002), whereas the genera Sphingomima (10 species) and Thenopa (four species) occur in Central and West Africa (Scoble, 1999). Sphingomima and Thenopa are externally similar, so the recovered sister-group relationship in the current phylogeny analysis was anticipated. In the current analysis, Hebdomophruda errans Prout, 1917 is isolated from other analyzed Hebdomophruda species (the others are included in Diptychini), highlighting the need for additional research. Krüger (1997, 1998) classified the genus Hebdomophruda into seven species groups on the basis of morphological characters, and H. errans group is one of them (Krüger, 1998). We do not describe a new genus for the taxon errans, nor do we combine it with any genus in the Drepanogynini, highlighting its uncertain taxonomic position (incertae sedis) pending more research. In the current analysis, Sphingomima discolucida Herbulot, 1995 is transferred from unassigned tribus combination to Drepanogynini, but as the type species of Sphingomima (S. heterodoxa Warren, 1899) was not analyzed, we do not transfer the entire genus Sphingomima into Drepanogynini. We highlight the uncertain taxonomic position of the taxon discolucida, acknowledging that it may eventually be included again in Sphingomima if the entire genus should be transferred to Drepanogynini.</p> <p>Diagnosis: Drepanogynini can be diagnosed by the combination of DNA data with up to 11 genetic markers (exemplar Drepanogynis mixtaria (Guenée, 1858)) ArgK (MK738841), COI (MK739615), EF1a (MK739960), IDH (MK740862), MDH (MK741181), Nex9 (MK741630), RpS5 (MK741991) and Wingless (MK742540). In the light of our phylogenetic results, the Drepanogynis group of genera, as classified earlier (Krüger, 2014), is split between two unrelated tribes (Drepanogynini and Diptychini). More research is needed to understand how other Drepanogynis species and the Drepanogynis group of genera sensu Krüger (1997, 1998, 1999, 2014) (at least 11 genera), should be classified.</p> <p>Boarmiini are the sister group to a clade that comprises Macariini, Cassymini, Abraxini and Eutoeini. We found that many species currently classified as Boarmiini are scattered throughout Ennominae. Boarmiini s.str. are strongly supported but are technically not monophyletic because of a large number of genera which need to be formally transferred fromothertribestoBoarmiini (G. Brehmetal., 2019, unpublisheddataforNeotropicaltaxa and L. Murillo-Ramos et al., 2019, unpublished data for other taxa). The results are principally in concordance with Jiang et al. (2017), who supported the monophyly of Boarmiini but with a smaller number of taxa.</p> <p>The divided valva in male genitalia was suggested as a synapomorphy of Macariini + Cassymini + Eutoeini by Holloway (1994). In addition, he proposed the inclusion of Abraxini in Cassymini. Although our findings support a close relationship, this group requires more study and a more extensive sampling effort. Similar findings were provided by Jiang et al. (2017) who suggested more extensive sampling to study the evolutionary relationships of these tribes.</p> </div>	http://treatment.plazi.org/id/038F87AD2B03FFA92EC55C40FCFE0F18	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B0DFFA42EC55ACFFA4F0CF7.text	038F87AD2B0DFFA42EC55ACFFA4F0CF7.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Archiearinae Fletcher 1953	<div><p>Archiearinae Fletcher, 1953</p> <p>The hypothesis presented in this study recovered Archiearinae as a monophyletic entity after some taxonomic rearrangements are performed. This subfamily was previously considered as sister to Geometrinae + Ennominae (Abraham et al., 2001), whereas Yamamoto &amp; Sota (2007) proposed them to be the sister-taxon to Orthostixinae + Desmobathrinae. Our findings agree with Sihvonen et al. (2011) who recovered Archiearinae as the sister-taxon to the rest of Geometridae excluding Sterrhinae and Larentiinae, although only one species was included in their study. Archiearis Hübner, (1823) is sister to Boudinotiana Esper, 1787 and these taxa in turn are sister to Leucobrephos Grote, 1874 (Fig. 4). The southern hemisphere Archiearinae require more attention. Young (2006) suggested that two Australian Archiearinae genera, Dirce and Acalyphes, actually belong to Ennominae. Our analyses clearly support this view and we therefore propose to formally transfer Dirce and Acalyphes to Ennominae (all formal taxonomic changes are provided in Table 2). Unfortunately, the South American Archiearinae genera Archiearides Fletcher, 1953 and Lachnocephala Fletcher, 1953, and Mexican Caenosynteles Dyar, 1912 (Pitkin &amp; Jenkins, 2004), could not be included in our analyses. These presumably diurnal taxa may only be superficially similar to northern hemisphere Archiearinae as was the case with Australian Dirce and Acalyphes.</p> </div>	http://treatment.plazi.org/id/038F87AD2B0DFFA42EC55ACFFA4F0CF7	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B0CFFA32EC55D3BFB460C2E.text	038F87AD2B0CFFA32EC55D3BFB460C2E.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Desmobathrinae Meyrick 1886	<div><p>Desmobathrinae Meyrick, 1886</p> <p>Taxa placed in Desmobathrinae were formerly recognized as Oenochrominae genera with slender appendages. Holloway (1996) revived Desmobathrinae from synonymy with Oenochrominae and divided it into the tribes Eumeleini and Desmobathrini.</p> <p>Desmobathrinae species have a pantropical distribution and they apparently (still) lack recognized morphological apomorphies (Holloway, 1996). Our phylogenetic analysis has questioned the monophyly of Desmobathrinae sensu Holloway because some species currently placed in Oenochrominae were embedded within the group (see also Sihvonen et al., 2011), and also the phylogenetic position of the tribe Eumeleini is unstable (see below). Desmobathrinae can be regarded as a monophyletic group after the transfer of Zanclopteryx, Nearcha and Racasta from Oenochrominae to Desmobathrinae, and the removal of Eumeleini (Table 2). Desmobathrinae as circumscribed here are an independent lineage that is sister to all Geometridae except Sterrhinae, Larentiinae and Archiearinae.</p> <p>The monobasic Eumeleini has had a dynamic taxonomic history: Eumelea was transferred from Oenochrominae s.l. to Desmobathrinae based on the pupal cremaster (Holloway, 1996), whereas Beljaev (2008) pointed out that Eumelea could be a member of Geometrinae based on the skeleto-muscular structure of the male genitalia. Molecular studies (Sihvonen et al., 2011, Ban et al., 2018) suggested that Eumelea was part of Oenochrominae s.str., but these findings were not well-supported and no formal taxonomic changes were proposed. Our analyses with IQTREE and RAxML recovered Eumeleini in two very different positions, either as sister to Geometrinae (SH-like = 93.6, UFBoot2 = 71) (Figs. 4 and 5), or as sister of Plutodes in Ennominae (RBS = 60) (Data S3). The examination of morphological details suggests that the position as sister to Geometrinae is more plausible: hindwing vein M2 is present and tubular; anal margin of the hindwing is elongated; and large coremata originate from the saccus (Holloway, 1994, our observations). The morphology of Eumelea is partly unusual, and for that reason we illustrate selected structures (Data S4), which include for instance the following: antennae and legs of both sexes are very long; forewing vein Sc (homology unclear) reaches wing margin; in male genitalia coremata are extremely large and branched; uncus is crossshaped (cruciform); tegumen is narrow and it extends ventrally beyond the point of articulation with vinculum; saccus arms are extremely long, looped; and vesica is with lateral rows of cornuti. However, the green geoverdin pigment concentration of Eumelea is low in comparison to Geometrinae (Cook et al., 1994). We tentatively conclude that Eumelea is probably indeed associated with Geometrinae. However, since eleven genetic markers were not sufficient to clarify the phylogenetic affinities of Eumelea, we provisionally place the genus as incertae sedis (Table 2).</p> </div>	http://treatment.plazi.org/id/038F87AD2B0CFFA32EC55D3BFB460C2E	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B0EFFA62EC55E53FCAA0B14.text	038F87AD2B0EFFA62EC55E53FCAA0B14.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Sterrhinae Meyrick 1892	<div><p>Sterrhinae Meyrick, 1892</p> <p>We included 74 Sterrhinae taxa in our analyses, with all tribes recognized in Forum Herbulot (2007) being represented. The recovered patterns generally agree with previous phylogenetic hypotheses of the subfamily (Sihvonen &amp; Kaila, 2004, Sihvonen et al., 2011). The genera Ergavia Walker, 1866, Ametris Guenée, 1858 and Macrotes Westwood, 1841, which currently are placed in Oenochrominae were found to form a well-defined lineage within Sterrhinae with strong support (SH-Like = 99 UFBoot2 = 100). These genera are distributed in the New World, whereas the range of true Oenochrominae is restricted to the Australian and Oriental Regions. Sihvonen et al. (2011) already found that Ergavia and Afrophyla Warren, 1895 belong to Sterrhinae and suggested more extensive analyses to clarify the position of these genera, which we did. Afrophyla was transferred to Sterrhinae by Sihvonen &amp; Staude (2011) and Ergavia, Ametris and Macrotes (plus Almodes Guenée, (1858)) will be transferred by P. Sihvonen et al. (2019, unpublished data).</p> <p>Cosymbiini, Timandrini, Rhodometrini and Lythriini are closely related as shown previously (Sihvonen &amp; Kaila, 2004; Õunap, Viidalepp &amp; Saarma, 2008; Sihvonenetal., 2011). Cosymbiini appear as sister to the Timandrini + Traminda Saalmüller, 1891 + Pseudosterrha Warren, 1888 and Rhodometrini + Lythriini clade. Lythriini are closely related to Rhodometrini as shown by Õunap, Viidalepp &amp; Saarma (2008) with both molecular and morphological data. Traminda (Timandrini) and Pseudosterrha (Cosymbiini) grouped together forming a lineage that is sister to the Rhodometrini + Lythriini clade (Fig. 2).</p> <p>Rhodostrophiini and Cyllopodini were recovered as polyphyletic with species of Cyllopodini clustering within Rhodostrophiini. Similar results were recovered previously (Sihvonen &amp; Kaila, 2004; Sihvonen et al., 2011), suggesting that additional work is needed to be done to clarify the status and systematic positions of these tribes. Sterrhini and Scopulini were recovered as sister taxa as proposed by Sihvonen &amp; Kaila (2004), Hausmann (2004), Õunap, Viidalepp &amp; Saarma (2008) and Sihvonenetal. (2011).</p> <p>Our new phylogenetic hypothesis constitutes a large step towards understanding the evolutionary relationships of the major lineages of Sterrhinae. Further taxonomic changes and more detailed interpretation of the clades will be dealt with by P. Sihvonen et al. (2019, unpublisheddata).</p> </div>	http://treatment.plazi.org/id/038F87AD2B0EFFA62EC55E53FCAA0B14	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
038F87AD2B11FFA62EC55B1CFD9B0C2F.text	038F87AD2B11FFA62EC55B1CFD9B0C2F.taxon	http://purl.org/dc/dcmitype/Text	http://rs.tdwg.org/ontology/voc/SPMInfoItems#GeneralDescription	text/html	en	Geometridae Leach 1815	<div><p>Geometridae Leach, 1815</p> <p>The phylogenetic hypothesis presented in this study is by far the most comprehensive to date in terms of the number of markers, sampled taxa and geographical coverage. In total, our sample includes 814 genera, thus representing 41% of the currently recognized Geometridae genera (Scoble &amp; Hausmann, 2007). Previous phylogenetic hypotheses were based mainly on the European fauna and many clades were ambiguously supported due to low taxon sampling. The general patterns of the phylogenetic relationships among the subfamilies recovered in our study largely agrees with previous hypotheses based on morphological characters and different sets of molecular markers (Holloway, 1997; Abraham et al., 2001; Yamamoto &amp; Sota, 2007; Sihvonen et al., 2011). However, the results of our larger dataset differ in many details and shed light on the phylogenetic relationships of several, poorly resolved, small subfamilies.</p> <p>Sterrhinae are recovered as the sister subfamily to the remaining Geometridae. This result is not in concordance with Sihvonen et al. (2011), Yamamoto &amp; Sota (2007) and Regier et al. (2009), who found a sister group relationship between Sterrhinae and Larentiinae which in turn were sister to the rest of Geometridae. Sihvonen et al. (2011) showed the Sterrhinae + Larentiinae sister relationship with low support, while Yamamoto &amp; Sota (2007) and Regier et al. (2009) included only a few samples in their analyses. Our analyses include representatives from almost all known tribes currently included in Sterrhinae and Larentiinae. The higher number of markers, improved methods of analysis, the broader taxon sampling as well as the stability of our results suggests that Sterrhinae are indeed the sister group to the remaining Geometridae. Sterrhinae (after transfer of Ergavia, Ametris and Macrotes, see details below), Larentiinae, Archiearinae, Geometrinae and Ennominae were highly supported as monophyletic. Oenochrominae and Desmobathrinae formed polyphyletic and paraphyletic assemblages, respectively.</p> <p>Table 2 Summary of formally proposed taxonomic changes.</p> <p>Transfer from Archiearinae to Ennominae</p> <p>Acalyphes Turner, 1926, to Ennominae: Diptychini</p> <p>Dirce Prout, 1910, to Ennominae: Diptychini</p> <p>Transfer from Oenochrominae to Desmobathrinae (Desmobathrini):</p> <p>Nearcha Guest, 1887</p> <p>Racasta Walker, 1861</p> <p>Zanclopteryx Herrich-Schäffer, 1855</p> <p>Transfer from Oenochrominae to Epidesmiinae:</p> <p>Abraxaphantes Warren, 1894</p> <p>Adeixis Warren 1987</p> <p>Dichromodes Guenée 1858</p> <p>Ecphyas Turner, 1929</p> <p>Epidesmia Duncan &amp; Westwood, 1841</p> <p>Phrixocomes Turner, 1930</p> <p>Phrataria Walker, 1863</p> <p>Systatica Turner, 1904</p> <p>New tribe combinations in Ennominae</p> <p>Psilocladia Warren, 1898, from unassigned to Gonodontini</p> <p>Oedicentra Warren, 1902, from Boarmiini to Gnophini</p> <p>Hypotephrina Janse, 1932, from unassigned to Gnophini</p> <p>Capusa Walker, 1857, from Nacophorini to Diptychini</p> <p>Mictodoca Meyrick, 1892, from Nacophorini to Diptychini</p> <p>Furcatrox McQuillan, 1996, from Nacophorini to Diptychini</p> <p>Amelora Guest, 1897, from Nacophorini to Diptychini</p> <p>Archephanes Turner, 1926, from Nacophorini to Diptychini</p> <p>Thalaina Walker, 1855, from Nacophorini to Diptychini</p> <p>Niceteria Turner, 1929, from Nacophorini to Diptychini</p> <p>Neazata Warren, 1906 from Caberini to Diptychini</p> <p>Idiodes Guenée, 1858 from unassigned to Diptychini</p> <p>Panhyperochia Krüger, 2013, from Nacophorini to Diptychini</p> <p>Mauna Walker, 1865, from Nacophorini to Diptychini</p> <p>Pareclipsis Warren, 1894, from unassigned to Diptychini</p> <p>Crambometra Prout, 1915, from unassigned to Diptychini Hebdomophruda Warren, 1897, from Nacophorini to Diptychini Pareclipsis Warren, 1894, from unassigned to Diptychini</p> <p>Capasa Walker 1866, from unassigned to Hypochrosini</p> <p>Omizodes Warren, 1894, from unassigned to Hypochrosini Metallospora Warren, 1905, from unassigned to Cassymini</p> <p>Obolcola Walker, 1862, from unassigned to Abraxini</p> <p>Chelotephrina Fletcher, 1958 from unassigned to Abraxini</p> <p>Cassephyra Holloway, 1994 from Cassymini to Abraxini</p> <p>Thenopa Walker, 1855 from unassigned to Drepanogynini Drepanogynis Guenée, 1858 from Nacophorini to Drepanogynini</p> <p>The monophylies of Oenochrominae and Desmobathrinae have long been questioned.</p> <p>Morphological studies addressing Oenochrominae or Desmobathrinae have been limited and the majority of genera have never been examined in depth. In addition, it has been very difficult to establish the boundaries of these subfamilies on the basis of morphological structures (Scoble &amp; Edwards, 1990). Sihvonen et al. (2011) showed that neither Oenochrominae nor Desmobathrinae were monophyletic, but these results were considered preliminary due to the limited number of sampled taxa, and as a consequence no formal transfers of taxa were proposed.</p> <p>The systematic status of Orthostixinae remains uncertain because it was not included in our study. Sihvonen et al. (2011) included the genus Naxa Walker, 1856, formally placed in Orthostixinae, and found it to be nested within Ennominae. However, only three genes were successfully sequenced from this taxon, and its position in the phylogenetic tree turned out to be highly unstable in our analyses. It was thus excluded from our dataset.</p> <p>Orthostixis Hübner, 1823, the type genus of the subfamily, needs to be included in future analyses.</p> </div>	http://treatment.plazi.org/id/038F87AD2B11FFA62EC55B1CFD9B0C2F	Public Domain	No known copyright restrictions apply. See Agosti, D., Egloff, W., 2009. Taxonomic information exchange and copyright: the Plazi approach. BMC Research Notes 2009, 2:53 for further explanation.		Plazi	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	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: 1-39, DOI: 10.7717/peerj.7386
