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Digitale Literatur/Digital Literature

Zeitschrift/Journal: Systematics and Phylogeny

Jahr/Year: 2019

Band/Volume: 77

Autor(en)/Author(s): Mally Richard, Hayden James E., Neinhuis Christoph, Jordal Bjarte H., Nuss Matthias

Artikel/Article: The phylogenetic systematics of and (: : ) inferred from DNA and morphology 141-204 77 (1): 141 – 204 2019

© Senckenberg Gesellschaft für Naturforschung, 2019.

The phylogenetic systematics of Spilomelinae and Pyraustinae (Lepidoptera: Pyraloidea: Crambidae) inferred from DNA and morphology

Richard Mally *, 1, James E. Hayden 2, Christoph Neinhuis 3, Bjarte H. Jordal 1 & Matthias Nuss 4

1 University Museum of Bergen, Natural History Collections, Realfagbygget, Allégaten 41, 5007 Bergen, Norway; Richard Mally [richard. [email protected], [email protected]], Bjarte H. Jordal [[email protected]] — 2 Department of Agriculture and Consumer Ser- vices, Division of Industry, 1911 SW 34th Street, Gainesville, FL 32608 USA; James E. Hayden [[email protected]] — 3 Technische Universität Dresden, Institut für Botanik, 01062 Dresden, Germany; Christoph Neinhuis [[email protected]] — 4 Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstraße 159, 01109 Dresden, Germany; Matthias Nuss [[email protected]] — * Corresponding author

Accepted on March 14, 2019. Published online at www.senckenberg.de/arthropod-systematics on May 17, 2019. Published in print on June 03, 2019. Editors in charge: Brian Wiegmann & Klaus-Dieter Klass.

Abstract. Spilomelinae and Pyraustinae form a species-rich monophylum of Crambidae (snout ). Morphological distinction of the two groups has been diffcult in the past, and the morphologically heterogenous Spilomelinae has not been broadly accepted as a natural group due to the lack of convincing apomorphies. In order to investigate potential apomorphic characters for Spilomelinae and Pyraustinae and to ex- amine alternative phylogenetic hypotheses, we conduct a phylogenetic analysis using 6 molecular markers and 114 morphological characters of the adults representing 77 genera of Spilomelinae and 18 genera of Pyraustinae. The results of the analysis of the combined data strongly suggest that Spilomelinae and Pyraustinae are each monophyletic and sister to each other. Wurthiinae is confrmed as ingroup of Spilomelinae, and Walker, 1859 as a non-spilomeline. Within Spilomelinae, several well supported clades are obtained, for which we propose a frst phylogeny-based tribal classifcation, using nine available and four new names: Hydririni Minet, 1982 stat.rev., Lineodini Amsel, 1956 stat.rev., Udeini trib.n., Wurthiini Roepke, 1916 stat.rev., Agroterini Acloque, 1897 stat.rev., Guenée, 1854 stat.rev. (= Siginae Hampson, 1918), Herpetogrammatini trib.n., Hymeniini Swinhoe, 1900 stat.rev., Asciodini trib.n., Trichaeini trib.n., Steniini Guenée, 1854 stat.rev., Nomophilini Kuznetzov & Stekolnikov, 1979 stat.rev. and Margaroniini Swinhoe & Cotes, 1889 stat.rev. (= Dichocrociinae Swinhoe, 1900; = Hapaliadae Swinhoe, 1890; = Margarodidae Guenée, 1854). The available name Syleptinae Swinhoe, 1900 could not be assigned to any of the recovered clades. Three tribes are recognized in Pyraustinae: Euclastini Popescu-Gorj & Constantinescu, 1977 stat.rev., Portentomorphini Amsel, 1956 stat.rev. and Pyraustini Meyrick, 1890 stat.rev. (= Botydes Blanchard, 1840; = Ennychites Duponchel, 1845). The taxonomic status of Tetridia Warren, 1890, found to be sister to all other investigated Pyraustinae, needs further investigation. The four Spilomelinae tribes that are sister to all other, ‘euspilomeline’ tribes share several plesiomorphies with Pyraustinae. We provide morphological synapomorphies and descriptions for Spilomelinae, Pyraustinae and the subgroups recognised therein. These characters allow the assignment of additional 125 genera to Spilomelinae tribes, and additional 56 genera to Pyraustinae tribes. New and revised combinations are proposed: Nonazochis Amsel, 1956 syn.n. of Guenée, 1854, with Conchylodes graph­ ialis (Schaus, 1912) comb.n.; Conchylodes octonalis (Zeller, 1873) comb.n. (from ); Hyperectis Meyrick, 1904 syn.n. of Hydriris Meyrick, 1885, with Hydriris dioctias (Meyick, 1904) comb.n., and Hydriris apicalis (Hampson, 1912) comb.n.; pandamalis (Walker, 1859) comb.n. (from ); Arthromastix pactolalis (Guenée, 1854) comb.n. (from ); coenostolalis (Hampson, 1899) comb.n. (from ); Prophantis xanthomeralis (Hampson, 1918) comb.n. (from Thliptoceras); Prophantis longicornalis (Mabille, 1900) comb.n. (from ); Charitoprepes apicipicta (Inoue, 1963) comb.n. (from Heterocnephes); Prenesta rubrocinctalis (Guenée, 1854) comb.n. (from ); Alytana calligrammalis (Mabille, 1879) comb.n. (from ). Epherema Snellen, 1892 stat.rev. with its type species E. abyssalis Snellen, 1892 comb.rev. is removed from synonymy with Syllepte Hübner, 1823. Ametrea Munroe, 1964 and Charitoprepes Warren, 1896 are transferred from Pyraustinae to Spilomelinae; Prooedema Hampson, 1891 from Spilomelinae to Pyraustinae; Aporocosmus Butler, 1886 from Spilomelinae to ; Orthoraphis Hampson, 1896 from Spi- lomelinae to Lathrotelinae; Hydropionea Hampson, 1917, Plantegumia Amsel, 1956 and Munroe’s (1995) “undescribed ex Boeo­ tarcha Meyrick” are transferred from Spilomelinae to .

Key words. Snout moths, phylogeny, tribal classifcation, morphology.

ISSN 1863-7221 (print) | eISSN 1864-8312 (online) | DOI: 10.26049/ASP77-1-2019-07 141 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

1. Introduction torium and the very reduced or absent gnathos, features common to both Spilomelinae and Pyraustinae, as due to parallelism. Furthermore, he considered none of the Pyraustinae and Spilomelinae comprise over 5,200 de- diagnostic features for Spilomelinae to be uniquely au- scribed species worldwide, accounting for about one third tapomorphic; instead, he diagnosed Spilomelinae by a of the species in Pyraloidea (Nuss et al. 2003 – 2019). It is combination of characters: chaetosemata absent, males estimated that about 50% of the pyraloid species are still without subcostal retinaculum, praecinctorium bilobed, undescribed on a global scale (Munroe 1972a), and Sut- tympanic frame protruding, spinulae distinctly tapered, ton et al. (2015) estimated that in Southeast 60% of male genitalia without well-developed gnathos, and fe- the species remain to be described. The knowledge about male genitalia without large rhombical signum. Solis eco­logy and especially host plant associations of the lar- & Maes’ (2003) cladistic study based on morphological vae is rather comprehensive for the species occurring in features of adults also implied that Pyraustinae and Spi- Europe (e.g. Hasenfuss 1960; Emmet 1988) and North lomelinae are not closely related. In contrast, a phyloge- America (e.g. Munroe 1972a,b, 1974b, 1976a). A review netic analysis of molecular data by Regier et al. (2012) of the known host plant data for the Oriental species has supported the monophyly of Pyraustinae + Spilomelinae; been given by Robinson et al. (2001). In recent years, rear- the diversity of both groups, however, was poorly sam- ing efforts of Lepidoptera caterpillars like those in Papua pled, with only two species of Pyraustinae and three spe- (Miller et al. 2007) and Costa Rica (Janzen cies of Spilomelinae included. Wurthiinae, characterised & Hallwachs 2009) collected from the local fora have by a number of morphological adaptations to their ant accumulated a large amount of host plant data for tropi- association, was recovered as ingroup of Spilomelinae. cal Spilomelinae. Altogether, spilomeline larvae feed on Recently, Lathrotelinae was revised and removed from a large variety of angiosperms, with varying degrees of Spilomelinae, comprising Diplopseustis Meyrick, 1884, host specifcity, and a few species feed on gymnosperms Diplopseustoides Guillermet, 2013, Lathroteles J.F.G. (e.g. Inoue & Yamanaka 2006) and ferns (e.g. Farah- Clarke, 1971 and Sufetula Walker, 1859 (Minet 2015). pour-Hagani et al. 2016). Larvae of Niphopyralis Hamp- The classifcation of Spilomelinae is confusing. The son, 1893 are associated with weaver ants, living in their subfamily includes 4,097 described species in 338 genera nests and feeding on ant larvae (Roepke 1916; Kemner (Nuss et al. 2003 – 2019). Many genera contain only a few 1923). Several species are known for their economic im- species, and 87 genera (26%) are monotypic. In contrast, pact on crops, among them the corn borers of the genus 20 genera comprise more than 50 species, collectively Hübner, 1825 (e.g. Nafus & Schreiner 1991), encompassing 50% of the species. The most species-rich the bean pod borer vitrata (Fabricius, 1787) (e.g. genera are Guenée, 1845, Hübner, 1808, Sharma 1998), the leafrollers of the genera Cnapha­ Glyphodes Guenée, 1854 and the heterogeneous genera locrocis Lederer, 1863 and Marasmia Lederer, 1863 (e.g. Syllepte Hübner, 1823 and Hübner, 1823. Pathak & Khan 1994) as well as the borers in Pyraustinae comprises 1,239 described species in 174 the genus Guenée, 1854 (e.g. Mally et al. genera, with 94 genera (52%) monotypic and only three 2015). Corn borers of the genus Ostrinia have become genera with more than 50 species: Hübner, model systems in basic and applied research, like for 1825, Hübner, 1823 and Schrank, 1802 population ecology, genetics and management as well (Nuss et al. 2003 – 2019). Tribes within Spilomelinae and as pheromone research (Elsworth et al. 1989; Burgio & Pyrau­ ­stinae have been proposed for recognition in the Mani 1995; Onstad & Gould 1998; Wang et al. 1998; past, but they usually served to segregate single genera Roelofs et al. 2002; Lassance 2010; Fuji et al. 2011). with aberrant morphology, e.g. the long-legged, narrow- Though there has been continuous progress in the winged Lineodini Amsel, 1956, Nomophilini Kuznetzov systematics of Spilomelinae and Pyraustinae, their cur- & Stekolnikov, 1979 and Hydririni Minet, 1982. There- rent classifcation is still largely based on typological fore, a comprehensive tribal classifcation has not been concepts. Spilomelinae and Pyraustinae are not easily thoroughly accepted. Munroe (1995) classifed the Neo- distinguishable based on external features and therefore tropical Spilomelinae into 15 genus groups plus many have long been considered as one taxon under the name unplaced genera, but he did not provide diagnoses for Pyraustidae, together with distinct groups like Schoeno­ these informal genus groups. biinae, , , Odontiinae and Gla­ The natural relationships among some Spilomelinae phy­riinae (Marion 1952). The distinction between Spi­ genera have been investigated (Sutrisno 2002a,b, 2003, lomelinae­ and Pyraustinae began to come into focus 2004, 2005, 2006; Sutrisno et al. 2006; Mally & Nuss through analyses of genitalia by Müller-Rutz (1929), 2010; Haines & Rubinoff 2012), but a large-scale phy- Pierce & Metcalfe (1938) and Marion (1952, 1954). logenetic analysis that takes the outstanding diversity of During the 1970s, the consensus was to classify Pyraus- Spilomelinae and Pyraustinae into account and identifes tinae into Spilomelini and Pyraustini (Munroe 1964, main lineages and their phylogenetic relationships has 1976a, 1995; Munroe & Solis 1998). In 1982, Minet not been published to date. split Spilomelinae from Pyraustinae, regarding them Our study provides the frst phylogenetic intra-sub­ only distantly related based on the lack of convincing family classifcation of Spilomelinae and Pyraustinae synapomorphies. He considered the bilobed praecinc- based on analysis of molecular, morphological and eco-

142 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019 GU829321 GU829302 GU829419 GU829411 GU829221 MK459853 MK459879 MK459880 MK459878 MK459871 MK459877 MK459869 MK459872 MK459870 MK459875 MK459876 MK459859 MK459860 MK459866 MK459867 MK459868 MK459873 MK459874 MK459856 MK459857 MK459861 MK459862 MK459863 MK459864 MK459865 MK459855 MK459858 MK459854 MK459851 MK459852 MK459849 MK459850 MK459848 EF-1a (1071bp) JX017948 GU829032 GU829014 GU829154 GU829143 GU828925 . (NCBI) in columns Origin and Collection MK459689 MK459677 MK459675 JF497027 JX017869 MK459696 MK459697 MK459695 MK459688 MK459694 MK459686 MK459692 MK459676 MK459683 MK459685 MK459673 MK459671 MK459669 MK459670 MK459667 MK459668 COI (1440bp) HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4 Peru ZSM MK506080 MK459674 Spain MTD MTD JF852437 Origin Collection Bolivia MTD MK506100 MK459679 Bolivia MTD MK506102 MK459672 Morocco MTD Udeini Guiana French MTD MK506103 MK459687 Tribus Steniini Steniini Steniini Asciodini Venezuela MHNG JF852400 Asciodini Guiana French R. Rougerie JN305177 MK459678 Asciodini Wurthiini Wurthiini Agroterini Germany MTD Agroterini ZMBN Spilomelini Cameroon A. Zwick Spilomelini MTD Nomophilini Guiana French MTD MK506091 MK459690 Nomophilini Bolivia MTD MK506106 MK459693 Nomophilini Leone Sierra Karisch T. Nomophilini Romania MTD Nomophilini Morocco MTD Nomophilini Bolivia MTD MK506088 MK459681 (MIDILINAE) Guiana French MTD Margaroniini China MTD JF852281 Margaroniini Malaysia ZMBN Margaroniini Malaysia ZMBN Margaroniini China MTD MK506074 MK459684 Margaroniini Cameroon A. Zwick Margaroniini Peru ZSM MK506101 MK459680 Margaroniini Peru ZSM MK506095 MK459682 Margaroniini Bolivia MTD MK506110 MK459691 Margaroniini Morocco MTD Margaroniini Peru ZSM Margaroniini UHIM ()(PYRALINAE) Germany (NCBI) MTD (NCBI) GU828590 GU828389 () (NCBI) (NCBI) GU828691 GU828487 (SCOPARIINAE) (NCBI) (NCBI) GU828709 GU828504 (ACENTROPINAE) (NCBI) (NCBI) GU828852 GU929816 (LATHROTELINAE) Germany MTD Species (Sepp, 1850) zebra Zeller, 1847 Zeller, fovealis Meyrick, 1888 iridia Meyrick, (Walker, 1859) (Walker, lauralis Walker, 1859 dariusalis Walker, (Cramer, 1777) tages (Cramer, cf. (Walker, 1859) diniasalis (Walker, designalis Guenée, 1854 (Walker, 1866) excelsalis (Walker, Zeller, 1850 uliginosellus Zeller, ebulealis (Guenée, 1854) (Hübner, 1825) (Hübner, pudoraria (Scopoli, 1863) nemoralis (Moore, 1867) (Moore, zebralis cf. (Linnaeus, 1758) farinalis pactolalis (Guenée, 1854) Munroe, 1970 guianensis Munroe, (Guenée, 1854) nervosalis Guenée, 1854 gordialis cf. (Walker, 1866) diminutalis (Walker, (Fabricius, 1775) punctalis (Fabricius, (Linnaeus, 1767) hyalinata (Zeller, 1852) (Zeller, cf inspersalis (Walker, 1859) pandamalis (Walker, (Walker, 1859) (Walker, perspectalis (Linnaeus, 1761) reticularis (Dyar, 1906) (Dyar, primordialis cf. (Walker, 1859) leodocusalis (Walker, (Guenée, 1854) fmbriauralis cf. medinalis (Guenée, 1854) cf. truncicolella (Stainton, 1849) (Guenée, 1854) favibasalis cf. cf. rufdiscalis Hampson, 1904 cf. Munroe, 1968 Munroe, novateutonialis catalaunalis (Duponchel, 1833) Munroe & Mutuura, 1968 & Mutuura, principis Munroe (Denis & Schifermüller, 1775) punctalis (Denis & Schifermüller, Arnia Pyralis Eporidia Aetholix Sufetula Synaphe Asciodes Cadarena Asturodes Apilocrocis Aristebulea Dichocrocis Agrioglypta Conogethes Clepsicosma Conchylodes Arthromastix Arthromastix 1775) gigantellus (Denis & Schifermüller, () (NCBI) (NCBI) GU828842 GU929806 ITBC058 MTD786 MTD649 MTD868 MTD776 MTD826 MTD557 MTD668 MTD882 MTD798 MTD797 MTD370 MTD393 MTD1316 MTD1041 MTD1047 MTD1357 MTD1061 MTD1281 MTD1319 MTD1323 MTD1331 MTD1325 MTD1019 MTD1347 MTD1329 MTD1354 MTD1016 MTD1328 MTD1307 MTD488A ZMBN097 ZMBN094 (WPH221) MM04967 MM03362 MM11223 MM09194 MM00051 DNA sample Genus refers to sequences obtained from GenBank, and neither origin nor storing collection are known. Table 1. Genetically investigated material with accession numbers. Second row of header states the primer pairs used to amplify respective genetic marker Table

143 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae — — GU829254 MK459914 MK459913 MK459909 MK459911 MK459912 MK459910 MK459877 MK459908 MK459907 MK459906 MK459882 MK459905 MK459881 MK459883 MK459887 MK459888 MK459891 MK459892 MK459893 MK459894 MK459895 MK459896 MK459897 MK459899 MK459901 MK459902 MK459903 MK459904 MK459884 MK459885 MK459886 MK459889 MK459890 MK459898 MK459900 EF-1a (1071bp) — — JX017965 JX017933 JX017926 GU828968 JF497031 JX017886 JX017856 JX017849 MK459725 MK459694 MK459724 MK459723 MK459722 MK459699 MK459721 MK459700 MK459704 MK459708 MK459717 MK459718 MK459719 MK459701 MK459702 MK459707 COI (1440bp) HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4 Peru ZSM MK506097 MK459726 Peru ZSM MK506081 MK459709 Origin Collection Bolivia MTD MK506112 MK459713 Bolivia MTD MK506090 MK459703 Angola MTD LN624711 MK459711 Angola MTD Morocco MTD Tribus Steniini Hydririni Guiana French MTD MK506082 MK459710 Hydririni Hydririni Lineodini Lineodini Lineodini Guiana French MTD MK506093 MK459720 Wurthiini Australia ANIC Agroterini Agroterini Bolivia MTD MK506092 MK459727 Agroterini (NCBI) (NCBI) GU828634 GU828432 Agroterini Philippines MTD Hymeniini SpilomeliniSpilomelini (NCBI) (NCBI) (NCBI) (NCBI) JX017856 JX017849 Nomophilini Morocco MTD Nomophilini Italy TLMF JF497031 MargaroniiniMargaroniini Guiana French Bolivia MTD MTD MK506083 MK506078 MK459728 MK459729 Margaroniini Morocco MTD Margaroniini Malaysia ZMBN MargaroniiniMargaroniini Costa Rica UHIM UHIM JX017886 Margaroniini Angola MTD Margaroniini Angola MTD Margaroniini Cameroon A. Zwick Margaroniini Bolivia MTD MK506087 MK459712 Margaroniini Peru ZSM Margaroniini Bolivia MTD MK506105 MK459714 Margaroniini Bolivia MTD MK506085 MK459715 Margaroniini Bolivia MTD MK506098 MK459716 Margaroniini Malaysia ZMBN Margaroniini Philippines MTD Margaroniini Yemen MTD Herpetogrammatini Bolivia MTD MK506099 MK459698 Herpetogrammatini Peru ZSM MK506075 MK459705 Herpetogrammatini Peru ZSM MK506096 MK459706 sp. sp. Species Dyar, 1913 vulnifca Dyar, (Rossi, 1794) vitrealis (Scopoli, 1763) ruralis Walker, 1859 sibillalis Walker, (Dyar, 1914) (Dyar, dissolvens Dognin, 1910 andrealis (Fabricius, 1787) (Fabricius, vitrata Hampson, 1896 pardalis (Fabricius, 1794) testalis (Fabricius, Guenée, 1854 humeralis (Walker, 1862) insolitalis (Walker, cf. stolalis Guenée, 1854 cf. cf. lygdamis Druce, 1902 cf. cf. pusialis Snellen, 1875 cf. (Moore, 1867) (Moore, zebralis cf. (Guenée, 1854) trapezalis chionesis Hampson, 1919 (Boisduval, 1833) (Boisduval, poeyalis cf. silicalis (Guenée, 1854) cf. (Fabricius, 1775) (Fabricius, derogata lutealis (Duponchel, 1833) (Walker, 1859) dorisalis (Walker, cf. (Walker, 1859) iphiclalis (Walker, cf. (Walker, 1859) hirtusalis (Walker, cf. (Hübner, 1796) (Hübner, perspectalis (Hübner, 1823) suppandalis (Hübner, africensis Mally et al., 2015 ornatalis (Duponchel, 1832) cf. obliqualis (Schaus, 1912) cf. (Guenée, 1854) phaeopteralis (Guenée, 1854) rubrocinctalis cf. (Wallengren, 1860) continuatalis (Wallengren, (Stoll in Cramer & Stoll, 1781) (Stoll in Cramer corope (Stoll in Cramer & Stoll, 1780) tedea (Stoll in Cramer cf. (Denis & Schifermüller, 1775) noctuella (Denis & Schifermüller, Yamanaka & Kirpichnikova, 1993 & Kirpichnikova, pseudocontortalis Yamanaka cf. Agroterini Malaysia ZMBN Palpita Patania Hydriris Maruca Liopasia Hileithia Omiodes Prenesta Prenesta Marasmia Marasmia Glyphodes Glyphodes Hodebertia Leucinodes Dichocrocis Gonocausta Obtusipalpis Niphopyralis Neoanalthes Lamprosema Leucochroma Ghesquierellana MTD784 MTD782 MTD994 MTD152 MTD820 MTD808 MTD364 MTD787 WPH197 WPH115 SDA008A MTD1018 MTD1324 MTD1033 MTD1342 MTD1483 MTD1318 MTD1321 MTD1566 MTD1285 MTD1337 MTD1004 MTD1043 MTD1562 MTD1349 MTD1251 MTD1284 MTD1341 MTD1340 MTD1046 MTD1283 MTD1282 MTD1565 ZMBN097 ZMBN104 ZMBN103 WPH252B MM00325 DNA sample Genus Table 1 continued. Table

144 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019 — GU829287 MK459939 MK459947 MK459948 MK459945 MK459946 MK459944 MK459941 MK459942 MK459943 MK459940 MK459938 MK459937 MK459936 MK459935 MK459934 MK459926 MK459931 MK459933 MK459927 MK459930 MK459928 MK459929 MK459925 MK459932 MK459923 MK459924 MK459922 MK459920 MK459921 MK459917 MK459918 MK459919 MK459916 MK459915 EF-1a (1071bp) JX017943 JX017930 GU829003 JF497033 LR135741 JX017826 JX017853 MK459744 MK459733 MK459759 MK459757 MK459756 MK459753 MK459755 MK459751 MK459750 MK459749 MK459748 MK459745 MK459746 MK459740 MK459738 MK459737 MK459732 MK459734 MK459731 COI (1440bp) HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4 MTD JF497033 NHMO Peru ZSM MK506076 MK459741 Peru ZSM MK506089 MK459743 Peru ZSM MK506084 MK459739 China MTD Kenya Kenya (NCBI) (NCBI) GU828675 GU828472 Origin Collection (NCBI) (NCBI) JX017853 Bolivia MTD LR135741 Canada MTD Morocco MTD JF852252 Tribus Udeini Udeini Udeini Hydririni Lineodini Venezuela MHNG JF852398 Trichaeini Euclastini Bulgaria S. Beshkov Euclastini Trichaeini Angola MTD Trichaeini Pyraustini Bolivia MTD MK506107 MK459760 Pyraustini Germany MTD Pyraustini Kyrgyzstan N. Pöll PyraustiniPyraustini Bolivia Germany MTD MTD MK506104 MK459754 Pyraustini Malaysia ZMBN PyraustiniPyraustini Bolivia Macedonia MTD MTD MK506108 MK459752 Pyraustini Romania MTD Pyraustini Bolivia MTD MK506109 MK459747 Pyraustini Agroterini Hymeniini Morocco MTD Spilomelini Spilomelini Spilomelini Guiana French MHNG Spilomelini Bolivia MTD MK506086 MK459735 Nomophilini Bolivia MTD MK506079 MK459736 Nomophilini USA FMNH Margaroniini Peru ZSM MK506111 MK459742 Margaroniini Angola MTD Margaroniini Philippines MTD Margaroniini Peru ZSM MK506094 MK459730 Portentomorphini Bolivia MTD MK506077 MK459758 Portentomorphini Australia UHIM JX017826 Portentomorphini Angola MTD sp. Species (Cramer, 1775) liris (Cramer, Poey, 1832 marialis Poey, Walker, 1859 Walker, salvalis Viette, 1957 gigantalis Viette, (Walker, 1859) (Walker, scyllalis (Walker, 1859) caritalis (Walker, (Hübner, 1796) nubilalis (Hübner, ecclesialis Guenée, 1854 (Hübner, 1796) (Hübner, ferrugalis (Hübner, 1796) (Hübner, pulveralis (Butler, 1878) (Butler, pantherata (Guenée, 1854) xanthialis Lederer, 1863 Lederer, multifarialis (Guenée, 1854) rantalis cf. hortulata (Linnaeus, 1758) (Hübner, 1796) aeruginalis (Hübner, (Fabricius, 1775) (Fabricius, recurvalis meticulosalis Guenée, 1854 (Fabricius, 1787) (Fabricius, perspicata cf. astrigalis Hampson, 1899 cf. muscosalis (Hampson, 1913) (Grote, 1882) (Grote, washingtonalis cf. taeniolalis (Guenée, 1854) cf. cf. minima (Hedemann, 1894) cf. cf. multiplicalis (Guenée, 1854) cf. (Hampson, 1918) xanthomeralis haemorrhoidalis (Guenée, 1854) Herrich-Schäfer, 1866 pilicornis Herrich-Schäfer, Sutrisno & Horak, 2003 Sutrisno & Horak, crenulata (Hampson, 1918) androstigmata cf. (Herrich-Schäfer, 1848) splendidalis (Herrich-Schäfer, (Stoll in Cramer & Stoll, 1781) (Stoll in Cramer forella (Stoll in Cramer & Stoll, 1782) phenice (Stoll in Cramer (Denis & Schifermüller, 1775) (Denis & Schifermüller, repandalis (Denis & Schifermüller, 1775) (Denis & Schifermüller, verbascalis Udea Udea Samea Anania Anania Syllepis Ostrinia Euclasta Trichaea Prenesta Udeoides Spilomela Syngamia Hyalorista Loxostege Oenobotys Cryptosara Prophantis Prophantis Rhimphalea Rhectosemia Pseudopyrausta Portentomorpha MTD477 MTD906 MTD388 MTD605 MTD553 MTD357 MTD956 MTD870 MTD783 MTD872 MTD774 MTD874 MTD650 WPH188 MTD276, MTD1344 MTD1322 MTD1343 MTD1350 MTD1327 MTD1466 MTD1558 MTD1484 MTD1017 MTD1338 MTD1315 MTD1247 MTD1320 MTD1467 MTD1248 MTD1358 MTD1235 MTD1015 ZMBN096 (WPH215) ZMBN011 ZMBN017 MM01851 DNA sample Genus Table 1 continued. Table

145 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae — — — — JX018024 HybRpS5r GU830697 GU830681 GU830800 GU830790 GU830604 MK460071 MK460070 MK460069 MK460068 MK460066 MK460067 MK460063 MK460064 MK460065 MK460058 MK460059 MK460062 MK460061 MK460060 MK460057 MK460055 MK460056 MK460053 MK460054 MK460052 MK459951 HybRpS5f/ RpS5 (576bp) MK459950 MK459949 EF-1a (1071bp) — — JX017919 JX017938 GU830095 GU830078 GU830230 GU830222 GU829979 MK459972 MK459973 MK459971 MK459970 MK459969 MK459968 MK459965 MK459966 MK459967 MK459959 MK459960 MK459961 MK459964 MK459962 MK459963 MK459957 MK459958 MK459955 MK459956 MK459953 MK459954 MK459952 HybIDHdegR HybIDHdeg27F/ — — — — — — — — — HybBurre JX017825 JX017861 MK460147 MK460143 HybFrigga/ HybFrigga/ MK459762 MK459761 COI (1440bp) — — HybCAD743f/ HybCAD743f/ HybCAD1028r HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4 Peru ZSM MK459771 China MTD MK459772 MK460142 Origin Collection (825bp) CAD GAPDH (654bp) IDH (657bp) Origin Collection Bolivia MTD MK459775 MK460145 Bolivia MTD MK459769 MK460141 Hawaii UHIM JX017825 Udeini Guiana French MTD MK459781 MK460149 Tribus Tribus Steniini Tetridiini Australia UHIM JX017861 Asciodini AsciodiniAsciodini Venezuela Guiana French R. Rougerie MHNG MK459774 MK460144 unplaced Wurthiini Wurthiini Agroterini Germany MTD MK459768 MK460140 Pyraustini Romania MTD Agroterini Malaysia ZMBN MK459766 MK460139 Pyraustini Germany MTD Spilomelini Philippines MTD MK459780 Nomophilini Leone Sierra Karisch T. — Nomophilini Bolivia MTD MK459777 Nomophilini Morocco MTD MK459773 (MIDILINAE) Guiana French MTD MK459764 MK460137 Margaroniini Malaysia ZMBN MK459782 MK460150 Margaroniini Cameroon A. Zwick MK459779 MK460148 Margaroniini China MTD Margaroniini Peru ZSM MK459776 Margaroniini Peru ZSM MK459778 MK460146 Margaroniini Morocco MTD MK459770 MargaroniiniMargaroniini Peru Australia ZSM UHIM MK459767 JX017793 (PYRALINAE) (NCBI) (NCBI) GU828092 GU829747 (PYRALINAE) Germany MTD MK459763 MK460136 (CRAMBINAE) (NCBI) (NCBI) GU828182 GU829811 (SCOPARIINAE) (NCBI) (NCBI) GU828197 GU829823 (ACENTROPINAE) (NCBI) (NCBI) GU828315 GU829906 (LATHROTELINAE) Germany MTD MK459765 MK460138 Species Species (Sepp, 1850) zebra Meyrick, 1888 iridia Meyrick, (Walker, 1859) (Walker, lauralis (Fabricius, 1794) (Fabricius, gilvata (Walker, 1859) diniasalis (Walker, designalis Guenée, 1854 (Walker, 1866) excelsalis (Walker, Zeller, 1850 uliginosellus Zeller, ebulealis (Guenée, 1854) (Hübner, 1825) (Hübner, pudoraria (Scopoli, 1863) nemoralis (Linnaeus, 1758) farinalis (Walker, 1859) (Walker, caletoralis pactolalis (Guenée, 1854) Munroe, 1970 guianensis Munroe, (Guenée, 1854) nervosalis Guenée, 1854 gordialis cf. (Walker, 1866) diminutalis (Walker, (Linnaeus, 1758) verticalis (Fabricius, 1775) punctalis (Fabricius, (Walker, 1859) pandamalis (Walker, (Zeller, 1852) (Zeller, inspersalis cf. (Dyar, 1906) (Dyar, primordialis cf. (Linnaeus, 1758) purpuralis (Guenée, 1854) fmbriauralis cf. medinalis (Guenée, 1854) cf. truncicolella (Stainton, 1849) (Guenée, 1854) favibasalis cf. cf. rufdiscalis Hampson, 1904 cf. Munroe, 1968 Munroe, novateutonialis catalaunalis (Duponchel, 1833) Munroe & Mutuura, 1968 & Mutuura, principis Munroe Arnia Midila Pyralis Tetridia Azochis Eudonia Aetholix Sufetula Agrotera Pyrausta Synaphe Asciodes Crambus Botyodes Cadarena Bocchoris Ategumia Asturodes Antigastra Apilocrocis Agathodes Anageshna Aristebulea Agrioglypta Conogethes Clepsicosma Conchylodes Arthromastix Arthromastix Schoenobius 1775) gigantellus (Denis & Schifermüller, (SCHOENOBIINAE) (NCBI) (NCBI) GU828306 GU829903 Cnaphalocrocis ITBC058 MTD826 MTD882 MTD798 MTD668 MTD776 MTD797 MTD370 MTD393 MTD560 MTD953 WPH054 MTD1041 MTD1319 MTD1281 MTD1019 MTD1347 MTD1329 MTD1331 MTD1325 MTD1061 MTD1354 MTD1016 MTD1328 MTD1307 MTD1326 MTD488A ZMBN094 (WPH221) (WPH209) MM04967 MM03362 MM11223 MM09194 MM00051 DNA sample Genus DNA sample Genus Table 1 continued. Table

146 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019 — — — — — — JX018009 JX018002 HybRpS5r MK460097 MK460095 MK460096 MK460093 MK460098 MK460092 MK460094 MK460090 MK460091 MK460089 MK460085 MK460086 MK460087 MK460088 MK460084 MK460080 MK460081 MK460082 MK460083 MK460078 MK460079 MK460101 MK460076 MK460077 MK460075 MK460100 MK460074 MK460099 MK460072 MK460073 HybRpS5f/ RpS5 (576bp) — — — MK460002 MK460000 MK460001 MK460003 MK459998 MK460004 MK459997 MK459999 MK459994 MK459995 MK459996 MK459989 MK459993 MK459988 MK459990 MK459991 MK459992 MK459987 MK459983 MK459984 MK459985 MK459986 MK459981 MK459982 MK460008 MK459979 MK459980 MK459978 MK460007 MK459974 MK459977 MK460005 MK460006 MK459975 MK459976 HybIDHdegR HybIDHdeg27F/ — — — — — — — — — — HybBurre MK460172 MK460175 MK460151 HybFrigga/ HybFrigga/ — — — HybCAD743f/ HybCAD743f/ HybCAD1028r Peru ZSM MK459802 MK460167 Spain MTD MK459788 Origin Collection (825bp) CAD GAPDH (654bp) IDH (657bp) Bolivia MTD MK459806 MK460170 Bolivia MTD MK459796 MK460162 Angola MTD MK459804 Angola MTD MK459801 MK460166 Morocco MTD MK459810 MK460174 Morocco MTD MK459789 MK460157 Tribus Steniini Steniini Steniini Hydririni Guiana French MTD MK459803 MK460168 Hydririni Hydririni Lineodini Lineodini Lineodini Guiana French MTD MK459812 MK460176 Wurthiini Australia ANIC MK459813 Agroterini Philippines MTD MK459797 MK460163 Hymeniini SpilomeliniSpilomelini (NCBI) (NCBI) (NCBI) (NCBI) JX017781 JX017777 Spilomelini Cameroon A. Zwick MK459790 Nomophilini Italy TLMF Nomophilini Bolivia MTD MK459786 MK460155 Nomophilini Morocco MTD MK459814 MK460177 NomophiliniNomophilini Guiana French MTD Romania MK459783 MTD MK460152 MK459785 MK460154 Margaroniini Malaysia ZMBN MK459811 Margaroniini Bolivia MTD MK459809 MK460173 Margaroniini Bolivia MTD MK459808 Margaroniini Bolivia MTD MK459807 MK460171 Margaroniini Bolivia MTD MK459805 MK460169 Margaroniini Yemen MTD MK459800 MK460165 Margaroniini Angola MTD MK459792 Margaroniini Philippines MTD MK459795 MK460161 MargaroniiniMargaroniini Cameroon Peru A. Zwick MK459793 ZSM MK460159 MK459794 MK460160 Margaroniini Malaysia ZMBN MK459787 MK460156 Margaroniini Angola MTD MK459815 MK460178 Margaroniini China MTD Margaroniini Bolivia MTD MK459784 MK460153 Herpetogrammatini Peru ZSM MK459798 Herpetogrammatini Peru ZSM MK459799 MK460164 Herpetogrammatini Bolivia MTD MK459791 MK460158 sp. sp. Species (Dyar, 1914) ens (Dyar, Dyar, 1913 vulnifca Dyar, Zeller, 1847 Zeller, fovealis Walker, 1859 sibillalis Walker, dissolv Dognin, 1910 andrealis Walker, 1859 dariusalis Walker, (Fabricius, 1787) (Fabricius, vitrata (Cramer, 1777) tages (Cramer, cf. Hampson, 1896 pardalis (Fabricius, 1794) testalis (Fabricius, (Walker, 1862) insolitalis (Walker, cf. stolalis Guenée, 1854 cf. cf. lygdamis Druce, 1902 cf. cf. pusialis Snellen, 1875 cf. (Moore, 1867) (Moore, zebralis cf. (Guenée, 1854) trapezalis chionesis Hampson, 1919 (Boisduval, 1833) (Boisduval, poeyalis (Fabricius, 1775) (Fabricius, derogata lutealis (Duponchel, 1833) (Walker, 1859) dorisalis (Walker, cf. (Linnaeus, 1767) hyalinata (Walker, 1859) (Walker, perspectalis (Linnaeus, 1761) reticularis (Walker, 1859) hirtusalis (Walker, cf. (Walker, 1859) leodocusalis (Walker, (Hübner, 1796) (Hübner, perspectalis (Hübner, 1823) suppandalis (Hübner, africensis Mally et al., 2015 ornatalis (Duponchel, 1832) cf. obliqualis (Schaus, 1912) cf. (Guenée, 1854) phaeopteralis (Stoll in Cramer & Stoll, 1781) (Stoll in Cramer corope (Denis & Schifermüller, 1775) punctalis (Denis & Schifermüller, (Denis & Schifermüller, 1775) noctuella (Denis & Schifermüller, Yamanaka & Kirpichnikova, 1993 & Kirpichnikova, pseudocontortalis Yamanaka cf. Agroterini Malaysia ZMBN Filodes Desmia Hydriris Maruca Mecyna Eporidia Metasia Liopasia Hileithia Nacoleia Hymenia Lineodes Cydalima Diasemia Megastes Marasmia Marasmia Diaphania Glyphodes Glyphodes Nomophila Hodebertia Leucinodes Haritalodes Dichocrocis Gonocausta Obtusipalpis Niphopyralis Duponchelia Neoanalthes Lamprosema Dolicharthria Diasemiopsis Leucochroma Eurrhyparodes Neoleucinodes Ghesquierellana Herpetogramma MTD787 MTD364 MTD808 MTD994 MTD820 MTD868 MTD786 MTD649 MTD782 MTD557 MTD152 WPH197 WPH115 MTD1340 MTD1341 MTD1046 MTD1284 MTD1562 MTD1251 MTD1043 MTD1349 MTD1337 MTD1282 MTD1565 MTD1004 MTD1283 MTD1566 MTD1318 MTD1285 MTD1316 MTD1321 MTD1483 MTD1357 MTD1047 MTD1323 ZMBN104 ZMBN103 ZMBN097 DNA sample Genus Table 1 continued. Table

147 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae — — — — — LR134887 JX018006 JX018019 JX018040 HybRpS5r GU830669 GU830638 MK460124 MK460122 MK460123 MK460121 MK460119 MK460120 MK460117 MK460118 MK460116 MK460114 MK460115 MK460113 MK460112 MK460111 MK460110 MK460109 MK460108 MK460107 MK460106 MK460105 MK460104 MK460103 MK460102 MK460125 HybRpS5f/ RpS5 (576bp) — LR134717 GU830062 GU830021 MK460037 MK460038 MK460035 MK460036 MK460034 MK460030 MK460031 MK460029 MK460032 MK460033 MK460027 MK460028 MK460026 MK460024 MK460025 MK460023 MK460022 MK460021 MK460019 MK460020 MK460018 MK460017 MK460016 MK460015 MK460014 MK460013 MK460012 MK460009 MK460039 MK460010 MK460011 HybIDHdegR HybIDHdeg27F/ — — — — — — — — — — HybBurre LR134626 MK460196 MK460190 MK460189 MK460188 MK460179 HybFrigga/ HybFrigga/ — — — — — — — — HybCAD743f/ HybCAD743f/ HybCAD1028r MTD MK459831 MK460197 NHMO MK459836 Peru ZSM MK459829 MK460194 Peru ZSM MK459828 MK460193 Peru ZSM MK459826 Peru ZSM MK459817 MK460182 NCBI NCBI GU828170 GU829798 China MTD MK459822 MK460186 Kenya Kenya (NCBI) (NCBI) Origin Collection (825bp) CAD GAPDH (654bp) IDH (657bp) Bolivia MTD LR134539 Canada MTD Morocco MTD MK459830 MK460195 Udeini Udeini Udeini Tribus Hydririni Lineodini Venezuela MHNG MK459823 MK460187 Trichaeini Euclastini Bulgaria S. Beshkov MK459837 Euclastini Trichaeini Angola MTD Trichaeini Pyraustini Romania MTD MK459834 MK460200 Pyraustini Pyraustini Bolivia MTD MK459833 MK460199 Agroterini Agroterini Bolivia MTD MK459818 MK460183 Agroterini Agroterini (NCBI) (NCBI) GU828133 GU829772 Hymeniini Morocco MTD MK459827 MK460192 Spilomelini Spilomelini Spilomelini Guiana French MHNG MK459825 Spilomelini Bolivia MTD Nomophilini USA FMNH MK459824 MK460191 Nomophilini Bolivia MTD Margaroniini Peru ZSM Margaroniini Angola MTD MK459832 MK460198 Margaroniini Philippines MTD Margaroniini Peru ZSM MK459821 MK460185 Margaroniini Bolivia MTD MK459820 Margaroniini Guiana French MTD MK459819 MK460184 Margaroniini Hawaii UHIM MargaroniiniMargaroniini Costa Rica Morocco UHIM MTD JX017808 MK459816 MK460180 MK460181 Portentomorphini Angola MTD MK459835 Portentomorphini Australia UHIM JX017788 Species (Cramer, 1775) liris (Cramer, Poey, 1832 marialis Poey, (Rossi, 1794) vitrealis (Scopoli, 1763) ruralis Viette, 1957 gigantalis Viette, (Walker, 1859) (Walker, scyllalis (Walker, 1859) caritalis (Walker, Guenée, 1854 humeralis ecclesialis Guenée, 1854 (Hübner, 1796) (Hübner, ferrugalis (Butler, 1878) (Butler, pantherata Lederer, 1863 Lederer, multifarialis cf. silicalis (Guenée, 1854) cf. (Guenée, 1854) rantalis cf. hortulata (Linnaeus, 1758) (Fabricius, 1775) (Fabricius, recurvalis (Walker, 1859) iphiclalis (Walker, cf. meticulosalis Guenée, 1854 (Fabricius, 1787) (Fabricius, perspicata cf. astrigalis Hampson, 1899 cf. muscosalis (Hampson, 1913) (Grote, 1882) (Grote, washingtonalis cf. multiplicalis (Guenée, 1854) cf. (Hampson, 1918) xanthomeralis haemorrhoidalis (Guenée, 1854) Herrich-Schäfer, 1866 pilicornis Herrich-Schäfer, (Guenée, 1854) rubrocinctalis cf. (Wallengren, 1860) continuatalis (Wallengren, Sutrisno & Horak, 2003 Sutrisno & Horak, crenulata (Hampson, 1918) androstigmata cf. (Herrich-Schäfer, 1848) splendidalis (Herrich-Schäfer, (Stoll in Cramer & Stoll, 1781) (Stoll in Cramer forella (Stoll in Cramer & Stoll, 1782) phenice (Stoll in Cramer (Stoll in Cramer & Stoll, 1780) tedea (Stoll in Cramer cf. (Denis & Schifermüller, 1775) (Denis & Schifermüller, verbascalis Siga Udea Udea Salbia Achyra Palpita Samea Samea alobathra Anania Anania Syllepis Patania Patania Terastia Phostria Euclasta Euclasta Trichaea Zebronia Omiodes Omiodes Prenesta Prenesta Prenesta Udeoides Spoladea Spilomela Syngamia Cryptosara Prophantis Prophantis Pycnarmon Rhimphalea Hy Rhectosemia MTD553 MTD870 MTD956 MTD357 MTD783 MTD872 MTD774 MTD874 MTD650 MTD784 WPH188 MTD276, SDA008A MTD1466 MTD1484 MTD1558 MTD1338 MTD1315 MTD1247 MTD1467 MTD1017 MTD1320 MTD1248 MTD1235 MTD1358 MTD1015 MTD1342 MTD1033 MTD1324 MTD1018 MTD1327 ZMBN011 ZMBN017 (WPH215) WPH252B MM01851 MM00325 DNA sample Genus Table 1 continued. Table

148 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

logical data of a global taxonomic sample. We also discuss the monophyly of the Neotropical genus groups proposed by Munroe (1995), since these represent the best recent attempt — to classify Spilomelinae. JX017996 JX018014 HybRpS5r MK460135 MK460134 MK460133 MK460131 MK460132 MK460127 MK460128 MK460130 MK460129 MK460126 HybRpS5f/ RpS5 (576bp)

2. Material and methods — MK460050 MK460051 MK460049 MK460047 MK460048 MK460045 MK460046 MK460041 MK460042 MK460044 MK460043 MK460040 HybIDHdegR

HybIDHdeg27F/ 2.1. Material

A broad range of Spilomelinae taxa was investigated to re- fect the morphological, ecological, evolutionary, and geo- — — — — — — graphical diversity of the group. 86 Spilomelinae species of HybBurre MK460204 HybFrigga/ HybFrigga/ 77 genera were studied, representing roughly one quarter of the genus-level diversity of Spilomelinae. In addition, we in- cluded 20 species of Pyraustinae representing 18 genera. We included 6 representatives of other Crambidae subfamilies as an internal outgroup taxon: (Stain- — ton, 1849) (Scopariinae), Zeller, 1850 HybCAD743f/ HybCAD743f/ HybCAD1028r (Crambinae), Schoenobius gigantellus (Denis & Schiffermül- ler, 1775) (Schoenobiinae), Midila guianensis Munroe, 1970 (Midilinae), Clepsicosma iridia Meyrick, 1888 (Acentropinae) and Sufetula diminutalis (Walker, 1866) (Lathrotelinae). The phylogeny is rooted with the external outgroup taxon consist- ing of the Synaphe punctalis (Fabricius, 1775) and Pyralis farinalis (Linnaeus,­ 1758) (both Pyralinae). See Table 1 for the list of taxa that were studied both genetically and Origin Collection (825bp) CAD GAPDH (654bp) IDH (657bp) Hawaiimorphologially. UHIM JX017770 Taxon sampling was primarily determined by the availability of freshly collected material suitable for the sequencing of the genetic markers of interest (see 2.2.1. Molecular methods). The studied taxa were identifed to ge- nus- or species level based on morphological investigations Tribus Tetridiini Australia UHIM JX017785 unplaced Pyraustini Romania MTD MK459847 MK460207 Pyraustini Germany MTD MK459846 MK460206 Pyraustini Germany MTD Pyraustini Bolivia MTD MK459845 MK460205 PyraustiniPyraustini Germany Kyrgyzstan MTD N. Pöll MK459841 MK459843 MK460203 Pyraustini Bolivia MTD MK459840 Pyraustini Malaysia ZMBN MK459842 MK460202 PyraustiniPyraustini Bolivia Macedonia MTD MTD MK459839 MK459838 MK460201 including genitalia dissection, as well as comparing the 5’ half Portentomorphini Bolivia MTD MK459844 of the mitochondrial COI gene sequence (‘DNA Barcode’) with the sequence data available on the Barcode of Life Da- tabase (BOLD, http://v4.boldsystems.org; Ratnasingham & Hebert 2007). For some taxa of interest, only one specimen was available for both molecular and morphological studies, resulting in the lack of the corresponding sex for investigation of its morphology. Where possible, we compensated for this

sp. lack by coding morphological features based on information Species from published literature (illustrations, descriptions). These

Walker, 1859 Walker, salvalis cases concerned the following taxa and consulted literature: (Fabricius, 1794) (Fabricius, gilvata (Hübner, 1796) nubilalis (Hübner, (Hübner, 1796) (Hübner, pulveralis (Walker, 1859) (Walker, caletoralis (Guenée, 1854) xanthialis (Linnaeus, 1758) verticalis (Hübner, 1796) aeruginalis (Hübner, (Linnaeus, 1758) purpuralis cf. taeniolalis (Guenée, 1854) cf. cf. minima (Hedemann, 1894) cf. Midila guianensis (Munroe 1970), Diasemiopsis leodocusalis (Walker, 1859) (Munroe 1957), Neoleucinodes dissolvens (Denis & Schifermüller, 1775) (Denis & Schifermüller, repandalis (Dyar, 1914) (Capps 1948), Euclasta gigantalis Viette, 1957 (Po­pescu-Gorj & Constantinescu 1977). Furthermore, we coded the morphology of closely related species for those taxa, where possible. The close relationship of those replacement specimens was evaluated by comparing Tetridia Pagyda Ostrinia Pyrausta Sitochroa Hyalorista Loxostege Uresiphita Paracorsia Oenobotys Psammotis the available genitalia, and where possible, COI barcode data. Pseudopyrausta Portentomorpha Those cases with replacement specimens are (male / female): An­ ageshna primordalis / A. cf. primordalis; Asciodes cf. gor­dia­ lis / A. quietalis; Azochis cf. rufdiscalis / A. rufdiscalis­ ; Me­- MTD560 MTD953 MTD477 MTD388 MTD906 MTD605

WPH054 gastes cf. pusialis / M. pusialis; Trichaea pilicornis / T. pro­ MTD1326 MTD1322 MTD1344 MTD1343 MTD1350 (WPH209) ZMBN096 DNA sample Genus

Table 1 continued. Table chyta; Hyalorista cf. taeniolalis / H. taeniolalis.

149 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

We state author and year of description of all genera subsequently amplifed in Sanger-sequencing PCR reac- and species at their frst mention in the text. Taxa identi- tions. Sequencing was done in both directions with the fed as ‘cf.’ have the author and year of description of the T7 and T3 primers, using the BigDye kit with this setup: closest species known to us. A list of investigated genita- 0.5 – 3.0 µl of PCR sample (depending on the sample’s lia slides is given in Supplement Table S1. band thickness on the agarose gel), 160 nM primer, 1 µl buffer, 0.5 µl BigDye, and adding up distilled water to 10 µl in total per reaction. Sequencing was conducted at 2.2. Methods the sequencing facilities of SNSD and UiB, Dept. of Mo- lecular Biology, or via Macrogen Europe. PCR, clean-up and sequencing PCR at SNSD was performed on a Mas- 2.2.1. Molecular methods tercycler ep gradient s (Eppendorf) or a PCR System 9700 (GeneAmp), at UiB a Bio-Rad 1000 thermal cycler was DNA extraction was done using the DNeasy Blood & used for PCR and sequencing PCR, and a MJ Research Tissue kit (Qiagen) or the NucleoSpin Tissue kit (Mach- PTC-200 thermal cycler for PCR clean-up. All sequences erey-Nagel) according to the manufacturers’ protocols. were proofread by eye and aligned manually using PhyDE The six genes COI, CAD, EF-1a, GAPDH, IDH and 0.9971 (Müller et al. 2008). All new sequence data have RpS5 were amplifed with the following primer pairs: been submitted to an open access nucleotide sequence COI in one large fragment with HybLCO (forward) database (GenBank; https://www.ncbi.nlm.nih.gov/gen- and HybPat (reverse) or as two shorter fragments with bank); accession numbers are compiled in Table 1. HybLCO (forward) and HybNancy (reverse) as well as HybJerry (forward) and HybPat (reverse); CAD with HybCAD743f (forward) and HybCAD1028r (reverse); 2.2.2. Morphological methods EF-1a (Elongation Factor 1-alpha) in one large frag- ment with HybOscar-6143 (forward) and HybEFrcM4 Genitalia were dissected according to Robinson (1976), (reverse) or as two shorter, overlapping fragments with with modifcations: The abdomen was cut open along one HybOscar-6143 (forward) and HybBosie-6144 (reverse) pleural membrane, cleaned, and embedded in medium as well as HybEF51.9 (forward) and HybEFrcM4 (re- under a cover slip to allow clear investigation of the tym- verse); GAPDH with HybFrigga (forward) and Hyb- panal organs. Female genitalia were stained with Chlo- Burre (reverse); IDH with HybIDHdeg27F (forward) razol Black. Male genitalia were either left unstained or and HybIDHdegR (reverse); RpS5 with HybRpS5f (for- were stained with Chlorazol Black or Eosin Y. ward) and HybRpS5r (reverse) (Wahlberg & Morphological structures were investigated using 2008; Haines & Rubinoff 2012). Each primer contains Leica M125 and M205C stereomicroscopes. Imagines a universal T7 (forward) or T3 (reverse) primer tail at were photographed with a Canon EOS 60D in combi- their 5’ end, which was used for sequencing (Wahlberg nation with a Canon EF 100mm 1:2,8 Macrolens and & Wheat 2008). Canon EOS Utility Version 2.10.2.0. A Leica CTR6000 All gene fragments were amplifed in 25 µl reactions. Microscope in combination with a Leica DFC420 camera The amplifcation protocol at the SNSD DNA lab was: and Leica Application Suite programme (Version 3.8.0) 200 nM of each primer, 200 µM dNTP mix, 2.5 µl Taq was used to photograph the genitalia. buffer, 1 mM MgCl2, 1 u BIO-X-ACT Short DNA Poly- Observed morphological features were coded accord- merase (Bioform), 2 µl DNA of concentration as extract- ing to the morphology character circumscriptions and ed, and distilled water added up to 25 µl in total per reac- compiled in a morphomatrix (Table 2) for all investigated tion. At the UiB DNA lab the PCR protocol was: 400nM taxa. Clepsicosma iridia (Acentropinae) was not studied of each primer, 800 µM dNTP mix, 2.5 µl Taq buffer morphologically, and is therefore omitted in Table 2. (incl. MgCl2), 0.75 u TaKaRa Ex Taq DNA Polymerase, 2 µl DNA of extracted concentration, and distilled water Morphological abbreviations in Figs. 3 – 15: an. – antrum; a.t. – added up to 25 µl in total per reaction. anal tube; ap.a. – apophysis anterioris; ap.p. – apophysis posterio- ris; apx. – appendix bursae; coe. – coecum of phallus; col. – col- The PCR programme for mitochondrial COI was: ini- liculum; cos. – costa; cos.b. – costa base; cos.d. – distal costa; cos. tial phase at 95°C for 5 min, 38 – 40 cycles at 95°C for ex. – dorsad process of basal costa; cos.p. – rod-shaped ventrad 30 s 50°C for 30 s and 72°C for 60 s, fnal phase at 72°C process of basal costa; crn. – cornutus or cornuti; c.b. – corpus for 10 min and cooling at 8°C. For the nuclear genes bursae; d.b. – ductus bursae; d.s. – ductus seminalis; div. – diver- CAD, EF-1a, GAPDH, IDH and RpS5 we ran a touch- ticulum; fo.ty. – fornix tympani; fb. – fbula; fa. – basal antennal fagellomeres; fr. – frons; fre. – frenulum; gna. – gnathos; h.p. – down PCR: 24 cycles at 95°C for 30 s 55°C with – 0.4°C hairpencil scerite(s); h.p.s. – hairpencil sclerites of the saccus; / cycle for 30 s and 72°C for 60 s + 2 s / cycle, then 12 hau. – haustellum; jx. – juxta; .p. – labial palps; lam. – lamella cycles at 95°C for 30 s 45°C for 30 s and 72°C for 120 s antevaginalis of ostium bursae; lob. – lobulus of lateral tympanal + 3 s / cycle, fnal phase at 72°C for 10 min and cooling case; mx.p. – maxillary palps; o.b. – ostium bursae; oc. – ocellus; at 8°C. p.a. – papilla anales; p.ph. – posterior phallus apodeme; ped. – pedicellus; pl.m. – pleural membranes; pl.tu. – pleural scale tufts PCR results were examined via gel electrophoresis of male abdominal segment 7; pl.sc. – pleural sclerites of male on a 1% agarose gel and GelRed as dying agent. Suc- abdominal segment 8; ret. – retinaculum; s2 – s8 – 2nd – 8th ab- cessful PCR samples were cleaned with ExoSAP and dominal sternite; sac. – sacculus; sac.d. – distal sacculus; sac. ex. –

150 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

extension of distal sacculus; sac.pr. – projection from central sac- for 30 Mio. generations, with sampling of every 1,000th culus; sc. – saccus of vinculum; sc.v. – ventral saccus tip; sca.f. – generation. The parameters for gamma shape, proportion raised scales on mesal side of fagellomeres; se.s. – sensillar setae of invariable sites, character state frequencies and GTR of fagellomeres; sig. – signum; t1 – t8 – 1st – 8th abdominal tergite; substitution rates were unlinked for the partitions, and the teg. – tegumen; teg.h. – hair-pencils on the dorsolateral tegumen; tran. – transtillum arm; tr.in. – transtillum inferior sensu Marion overall rate was allowed to vary across partitions. The ini- 1954; ty.fr. – tympanic frame; unc. – uncus; u.ch. – uncus chaetae; tial 25% of the were discarded as burn-in. Effective v.s. – venula secunda; v.va. – ventral valva edge; ves. – vesica; sampling sizes (ESS) and the degree of convergence of vin. – vinculum; vin.d. – dorsal joint of vinculum with valva. the runs were evaluated in Tracer (Rambaut et al. 2014). The phylogenetic trees were annotated using TreeGraph 2.14.0-771 beta (Stöver & Müller 2010). 2.2.3. Phylogenetic analysis In addition, a Maximum Likelihood (ML) analysis of the gene-partitioned molecular dataset was done using For the phylogenetic analyses, the nucleotide sequences RAxML-HPC2 (8.2.10) (Stamatakis 2014) on XSEDE of the genetic data were used. Initial Maximum Likeli- through the CIPRES V 3.3 online platform (Miller et al. hood analyses showed that analysis of the amino acid 2010). sequences translated from the nucleotide data resulted in WinClada 1.00.08 (Nixon 2002) was used to derive poorly resolved topologies with branch supports mostly ancestral morphological characters from the topology >> 50 percent; amino acid sequences were therefore dis- that was observed in the majority of analyses. Unambig- regarded as phylogenetic data source. uous synapomorphies, and in addition those derived from The sequence data were investigated for potential slow optimization (slow optimization or DELTRAN, substitution saturation in third codon positions (Xia et al. Swofford & Maddison 1987) are plotted on the topology 2003; Xia & Lemey 2009) using DAMBE5 (Xia 2013). and summarized. Apomorphies are included in the diag- RogueNaRok (Aberer et al. 2013) was used to screen the noses of clades (see Phylogenetic classifcation section) molecular data for rogue taxa, which were subsequently except if they are very homoplastic in that clade. Char- excluded from the analysis. acters were mapped on the consensus of the parsimony We analysed the concatenated molecular and mor- cladograms as well as the Baysian trees for the sake of phological data with two different partitioning schemes: methodological consistency (Assis 2015). GENES and TIGER. In the GENES scheme, we placed A parsimony analysis was conducted with TNT 1.5 each gene and the morphological data into a separate (Goloboff & Catalano 2016). All states were non-addi- partition, resulting in 7 partitions. In the TIGER scheme, tive and equally weighted, and gaps were treated as miss- following Rota & Wahlberg (2012), we partitioned the ing data. A traditional search plus the parsimony ratchet molecular dataset in terms of evolutionary site rates using (Nixon 1999) and branch-swapping was done (com- the programme TIGER (Cummins & McInerney 2011). mands: mxram 100; cc-.; collapse [; rs 1; hold 10000; rat: We chose initial partitioning into 10 bins and pooled the iter 50; mu: hold 20 replic 100 rat; bb;). Ratchet com- bins with < 100 sites with bin 1, resulting in 5 molecular mands were the default values: stop when 14 substitu- partitions, with bin 1 (incl. bins 2 – 6, each with < 100 tions made, 4% upweight and downweight probability, sites) = 1798 sites, bin 7 = 182 sites, bin 8 = 684 sites, bin 50 total iterations, alternating equal weights. One hun- 9 = 1291 sites, and bin 10 = 1213 sites; morphology was dred replications were done, saving 20 trees per repli- treated as a separate 6th partition. cation. In addition to equal weights, implied weighting We used jModeltest v2.1.4 (Guindon & Gascuel 2003; (Goloboff 1993) was explored under a range of k-pa- Darriba et al. 2012) to infer the models that best refect rameter values in TNT with the same search parameters. the sequence evolution of the genetic data. The resulting To try to resolve incongruence among cladograms, we models for the GENES partitioning are: TIM3+G+I mod- ran the IterPCR script provided by Pol & Escapa (2009). el for the COI partition, SYM+G+I model for the EF-1a This script suggests characters to recode, which is not partition, TVM+G+I model for the GAPDH partition, and done by the application embedded in TNT 1.5. GTR+G+I model for the CAD, IDH and RpS5 partitions. The TIM3 model is not implemented in MrBayes, and for this and the TVM model we used the GTR model instead. We omitted the invariant sites (I) parameter from the mod- 3. Results els since the parameters G and I are strongly correlated, and fewer parameters improved the analysis time (Sul- livan & Swofford 2001). The TIGER partitioning scheme 3.1. Molecular data was analysed under the GTR+G model. For the morpho- logical partition, we applied the Mk model with gamma rate variation (Lewis 2001). The concatenated dataset We present new genetic data for 100 taxa. In addition, was analysed with MrBayes version 3.2.6 (Ronquist et we complemented the genetic data for four taxa from al. 2012) on the CIPRES online platform (Miller et al. the study of Haines & Rubinoff (2012), for which we 2010) using Extreme Science and Engineering Discovery obtained the original DNA extracts from Will Haines Environment (XSEDE). Two parallel runs were set up (University of Hawaii): vouchers WPH209, WPH215,

151 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

WPH221, and WPH252. COI sequencing was successful 3.2. Morphological data for all samples except for the 3’ half of ‘Thliptoceras’ xanthomeralis Hampson, 1918 (DNA voucher ZMBN Lep017). Sequencing success for CAD was 85% (of n = Morphological data was coded from investigation of dried 100 samples), for the 1st part of EF-1a 94% (n = 100), adult specimens and their genitalia. For a complete list of for the scond part of EF-1a 100% (n = 104), for GAPDH genitalia slides of species investigated in the context of 70% (n = 104), for IDH 97% (n = 104), and for RpS5 85% this study (beyond the taxa included in the phylogenetic (n = 100). The amplifcation of RpS5 with the primers dataset), see Electronic Supplement File 1. of Wahlberg & Wheat (2008) failed for all taxa in the The morphological investigation resulted in the recog- tribes Udeini and Lineodini (see for tribes) nition of 115 variable characters for all 114 taxa. Of these except for 37% of the sequence length of Lineodes vul­ characters, 91 are binary, and 24 are multistate. Nineteen nifca, probably due to a lack of match between primer characters code features of the head and thorax including and attachment sequence. legs and wings, 23 of the abdomen including the tym- The molecular alignment has a length of 5,223 base- panal organs but excluding the genitalia, 47 of the male pairs (bp), with 1,440 bp accounting for COI, 825 bp for genitalia, 25 of the female genitalia and one character of CAD, 1,071 bp for EF-1a, 654 bp for GAPDH, 657 bp the locality of larval feeding. Character 115 (locality of for IDH, and 576 bp for RpS5. larval feeding) was coded from literature data, and the The CAD sequence of (GenBank following literature was used: Hinckley (1964), Genty & accession no. MK459834) lacks three codons (9 bp, i.e. Mariau (1975), Munroe (1976), Allyson (1984), Com- three amino acids in the respective protein product) com- mon (1990), Nuss (2005), Speidel (2005), Slamka (2008, pared to all other CAD sequences incorporated in our 2013), Hayden et al. (2013), Leraut (2014) and Pereira dataset. These three codons are present in the congeneric et al. (2014), as well as a personal observation of Leuci­ species, A. hortulata, and they code for the amino acids nodes africensis Mally et al., 2015 from Marja van der Isoleucine-Alanine-Valine. This three-codon deletion is Straten (pers. comm.). situated in a variable region of the CAD gene, where es- The morphomatrix is shown in Table 2. The defni- pecially the second codon is coding for a variety of dif- tions of the morphological characters and their states are ferent amino acids among the investigated taxa. A three- as follows: codon deletion at the identical location in the CAD se- 1 Presence of anteriad-directed projection medially on quence was observed in other Pyraustinae taxa believed frons: (0) absent (Fig. 11C); (1) present (Fig. 4B). to be closely related to Anania (Kai Chen, pers. comm.), 2 Presence of haustellum: (0) absent (Fig. 6A; Roepke so that this deletion might represent a synapomorphy for 1916: fg. 2); (1) present (Fig. 4B). these taxa. 3 Presence of transverse rim on anterior or mesal face The long terminal branch of Niphopyralis chionesis of pedicellus [male]: (0) absent (Fig. 8F); (1) present Hampson, 1919 in the phylogenetic results of Mutanen (Fig. 11C). et al. (2010) and Regier et al. (2012) is suspicious, and we therefore re-sequenced this species from available 4 Presence of a crest or prong of raised scales on mesal material (voucher no. MTD152) to evaluate whether the side of fagellomeres [male]: (0) absent (Fig. 11C); data from the previous studies might be compromised. (1) present at ca. 1/3 of antenna length, crest forming Our resulting sequence coverage was comparable to that a triangular prong proximally (Fig. 11D); (2) present in proximal part of antenna (Fig. 8F). of Mutanen et al. (2010), where the frst half of EF-1a and the entirety of the GAPDH were unsuccessful in 5 Length of sensillar setae at basal antennomeres rela- sequencing, just as in our results. Our sequenced data tive to diameter of basal antennomeres [male]: (0) ≤ largely matches that of the Mutanen et al. (2010) dataset, 50% (♂ in Fig. 10I, Fig. 11C); (1) > 50% (♂ in Fig. with only a few nucleotide differences between the cor- 10H, Fig. 11D). responding sequences of the two investigated specimens. 6 Length of cilia at antenna base in female compared This result strongly suggests that the long terminal branch to male: (0) of equal length (Fig. 10I); (1) shorter of N. chionesis in former studies is not due to sequence (Fig. 10H). contamination. However, since the same DNA lab proto- 7 Presence of ocelli: (0) absent (Fig. 6B); (1) present col (Wahlberg & Wheat 2008) was used in the Mutanen (Fig. 11C). et al. (2010) study as well as in the present study, the 8 Direction of 3rd labial palpomere: (0) dorsal (Fig. sequencing of pseudogenes cannot be ruled out, although 11C); (1) porrect (Fig. 4B). no reading frame shifts or stop codons occur in any of 9 Intersexual size differences of 3rd labial palpomere: the investigated N. chionesis sequences, suggesting that (0) well developed in both sexes (Maes 1995: pl. 5); they code for functional proteins. None of the nucleotide (1) short in the male (Fig. 4B); (2) short in both sexes sequences of N. chionesis is found to be exceptionally (Fig. 6A; Roepke 1916: fg. 3). divergent from those of other investigated taxa, and ob- 10 Length of maxillary palpi: (0) long enough to hypo- served nucleotide substitutions relative to the other taxa thetically come in contact with each other (Fig. 11C); mostly result in synonymous amino acid codons, i.e. they (1) minute to obsolete, cannot hypothetically get in encode the same amino acid. contact with each other (Fig. 6A).

152 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

11 Presence of broad scale tuft on distal foreleg tibia segment 5, one small scale tuft on anterior ends of (not to be confused with epiphysis): (0) absent; (1) segments 6 and 7: (0) absent (Fig. 12G); (1) present present. (Fig. 13C). 12 Presence of tuft of long scales on distal foreleg fe- 29 Presence of large, oval pleural scale tufts on each mur: (0) absent; (1) present. side of male abdominal segment 7, with a presum- 13 Presence of a longitudinal groove bearing a hair pen- ably secretory opening in its anterior centre: (0) ab- cil on male midleg tibia: (0) absent (Frolov et al. sent (Fig. 3H); (1) present (Fig. 14F). 2007: fg. 1A); (1) present (Frolov et al. 2007: fg. 30 Outline of central anterior edge of male sternite 7: 1B,C). (0) straight to slightly undulate (Fig. 6F); (1) anteri- 14 Presence of tibio-abdominal scale brush [male]: lon­ orly projecting protuberance or spine (Fig. 3H); (2) gitudinal line of thin, spatulate scales on male hind­ arch-shaped recession (Fig. 8G). leg’s proximal tibia in contact with an area of ventrad scales on pleural membranes of abdominal segment 31 Outline of central posterior edge of male sternite 7: (0) straight (Fig. 8H); (1) with pair of posteriad, 2: (0) absent; (1) present (Mey & Speidel 2010: fgs. 5, 10). curved spines running dorsally of sternite 8 (Fig. 15 Number of apical spurs on hindtibia: (0) 4, a proxi- 6F); (2) with pair of posteriad lobes (Fig. 15E); (3) a mal and a distal pair (Fig. 10F, G); (1) 2, only a dis- wide V-shaped recession (Fig. 8E). tal pair (as in fg. 10F,G, but without proximal spur 32 Outline of anterior edge of male tergite 8: (0) straight pair). to convex (Fig. 6F); (1) with triangular and straight- 16 Length of metatibial proximal inner spur relative to edged or semicircular indentation (Fig. 3H). tibial segment between this and the distal spur pair: 33 Sclerotization of male tergite 8: (0) homogenous (0) < 1/2 (distance “d”) (Fig. 10F); (1) ≥ 1/2 (dis- (Fig. 9D); (1) heterogenous, i.e. with distinct sclero­ tance “d”) (Fig. 10G). ti­zation pattern (Fig. 3H). 17 Presence of feld of enlarged, raised scales on male 34 Sclerotization pattern on male tergite 8: (0) central central forewing costa: (0) absent (Fig. 7A); (1) pre- longitudinal strip; (1) longitudinal strip, bifurcating sent (Fig. 9A). anteriorly into a Y-shape (Fig. 3H). 18 Form of retinaculum at costal base of forewing un- 35 Presence of a feld of setose scales on the anterior derside in males: (0) simple brush of straight hairs ends of the male tergite 8’s Y-shaped sclerotisation: (Fig. 8I); (1) cuticle protruded as a retinacular hook (0) absent (Fig. 8H); (1) present (Fig. 11F). (frenulum hook sensu Forbes 1926: fg. 7; Popescu- 36 Presence of U-shaped sclerotisation on lateral and Gorj & Constantinescu 1977: fg. 3b). anterior edge of male sternite 8: (0) absent (Fig. 9D); 19 Number of frenulum bristles in female: (0) one (Fig. (1) present (Fig. 11F). 4A); (1) two (Popescu-Gorj & Constantinescu 1977: fg. 3a). 37 Presence of an anterolaterad sclerotized lobe on each 20 Splitting of praecinctorium: (0) strong (Marion 1954: side of anterior edge of male sternite 8, running dor- fg. 2); (1) weak to absent (Marion 1954: fg. 1). sad of sternite 7: (0) absent (Fig. 8H); (1) present 21 Presence of lobulus on lateral edge of tympanal case: (Fig. 6G). (0) absent (Fig. 6C); (1) present (Fig. 8A). 38 Presence of central hair scale patch(es) on anterior 22 Shape of fornix tympani surface: (0) projecting from edge of male sternite 8: (0) absent (Fig. 8H); (1) pre- the tympanic frame (Minet 1983: “cd.” in fg. 30; sent (Fig. 3H). Maes 1985: “f.ty.” in pls. 1A, 1D); (1) recessed with- 39 Presence of median U-shaped recession or deep in the frame (Maes 1985: pl. 1E, “f.ty.” in pl. 2C). notch on posterior edge of male sternite 8: (0) absent 23 Direction of fornix tympani projection: (0) ven- (Fig. 3H); (1) present (Fig. 6G). tral (Minet 1983: “cd.” in fg. 30); (1) lateral (see 40 Presence of a sclerite on each pleural membrane of Hayden 2013: fgs. 18, 19, 21, 22). male segment 8: (0) absent (Fig. 6F); (1) present 24 Presence of venulae secundae: (0) absent (Fig. 6C); (Figs 3H, 9D). (1) present (Fig. 8A). 41 Shape of pleural sclerite on male segment 8: (0) slim 25 Course of venulae secundae: (0) converging (Fig. longitudinal strip (Fig. 9D); (1) broad semicircle 5C); (1) in posterior half parallel or diverging (Fig. (Fig. 3H). 8A). 42 Presence of a feld of setae anterior on pleurites of 26 Presence of lateral anteriad lobe on each side of ante- rior edge of male sternite 3: (0) absent (Fig. 6C); (1) male segment 8: (0) absent (Fig. 3H); (1) present. present (Fig. 7G). 43 Presence of uncus: (0) absent (reduced) (Fig. 3A); 27 Presence of pleural scale tufts on each side of the (1) present (Fig. 3D). male abdomen, one small scale tuft anteriorly on 44 Shape of uncus: (0) conical, non-capitate (Fig. 3D); segment 6 and one large scale tuft anteriorly on seg- (1) capitate (Fig. 3G). ment 7: (0) absent (Fig. 3H); (1) present (Fig. 8H). 45 Shape of apical uncus: (0) single head (Fig. 3D); 28 Presence of pleural scale tufts on each side of the (1) bi- or trifurcate head (Fig. 9E); (2) two separate male abdomen, one large scale tuft anteriorly on heads (Figs. 9G, 11E).

153 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111 0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345 0100111100 000001010? 01?1000000 000?000000 ?010010011 2000010000 ??000????0 00?000010? 10011001B0 1000?0000? 010110???? 000?0 0100111000 000001010? 01?0?00000 000?000000 ?010010411 0000200000 ??000????0 00?000010? 1001100100 000100000? 0000?0???? 000?E 0100001100 00000100?0 1010?00000 001??00000 ?011010421 ???0?????0 ??000????0 01?1000000 10011011B2 0111000110 0000?11100 000?1 0100011100 0000010010 1011000000 000??00000 ?011010001 ???0000000 ??000????0 00?0000000 1000?000?0 1001000010 0000?1100? 00120 01????10?0 ??????0??? ?????????? ?????????? ??10010?01 ???0000000 ??000????0 00?0000000 10011?010? ?????????? ?????????? ????? 0000111100 0000010000 1011000000 010??10010 ?010010401 ???0000000 ??000????0 00?0000100 10011001B0 1000?00010 1000?0???? 000?2 01000?00?0 00000100?1 0011100000 000??00000 ?00??0???? ??0??00000 ??000????0 00?0000200 10011001D2 100110000? 000110???? 000?4 0100111001 0000010010 0001100000 0111010000 ?011110312 4100001001 0000111100 0000000100 1000?101D2 000110000? 0010?0???? 000?? 0100001121 0000010010 1001100000 0111010001 001000??10 2100000011 0000122100 0110100200 1000?10120 0001100011 0000?10??? 0011C 0100011120 0000010010 0001100000 0111010001 0110010301 2100110111 10010200?0 0110100100 1000?101B1 000110000? 0001011100 000?0 0100111000 0000010010 0000?00000 000??00000 ?010010310 1100120100 ??001200?0 0011000100 10110111B0 001100000? 0100?1100? 000?0 0100111000 0000010000 1000?00000 0111010001 0011011112 0000000011 ??000????0 00?0000200 1000?10121 000010000? 0200?0???? 000?4 0100001120 00?000001? 1000?00000 010??00000 ?01101101? 0000010000 ??00100100 0111100200 1000?10122 0001000011 0010?0???? 000?C 0100111121 0000010010 0??0?00000 000??00000 ?01001A002 0000200000 ??00110100 0010000000 100110012? ?????????? ?????????? ????? 0100011100 0000010010 0000?00000 000??10000 ?011011002 0000200000 ??00110100 0110000110 1001000112 001100001? 0011011??? 000?? 0100111120 0000000010 0001100000 000??10001 0010011012 000010011? ??00100100 0101000100 1011010110 0001000010 0000?1000? 000?? 0101111020 0100010010 0000?00000 0111010000 ?011011012 1100020001 1101010100 0121100200 1000?00100 000111000? 0011010??? 000?? 01001?10?0 00000100?0 0001100000 0111110000 ?011111012 1100?00101 1100132100 0121001100 1000?001B0 001110000? 0010?10??? 000?? 0101111021 1100010010 0000?00000 0111110001 0011211012 2100120001 1100122100 0121000110 1000?00120 0011?0000? 001??10??? 000?0 0100001021 0000010010 0000?00000 0111010001 011101120? 2000000111 0001022100 0111100200 1001010120 000110000? 0010?10??? 000?? 0100001120 00000100?0 0000?00000 0011010001 000??0???2 1000020101 00001220?0 0111001210 1001010112 000110000? 000111105? 000?? 01021?11?1 00?00100?0 0001100000 4011010001 0011011011 2000000001 11010230?0 0000100200 1000?101D0 100110000? 0001011122 000?? 0100111120 0000010010 0001000000 0111010000 ?00??0???? 0000110011 01000????0 00?0001200 10010111D? 1001?1001? 001101A00? 000?3 0100001120 0000010010 1001100000 0010010001 0011011012 2100020011 11000????0 00?1100100 1000?10121 000110000? 001100???? 000?0 0100001120 0000010010 0001100000 0111010000 ?011011222 0000?10111 00001200?0 0110100200 0000?101B0 000110000? 0?00?10??? 0010? 0100011120 0000011010 1000?00000 0111010001 001121101? 0010001101 0001020100 0001010100 1000?00120 0001000010 001101100? 000?0 0100001021 0000000010 0000?00000 010??10000 ?0100101?? 0000120000 ??00120100 0110010100 1001101120 0001000011 000101102? 0012? 01000?10?1 00000100?0 0001100001 0111010001 0011010012 2000000001 0101000100 0111110200 1000?101B? ?????????? ?????????? ????? 0100001120 0000000010 1001100001 0?11010001 001111A202 2100100111 0100122100 0111100200 1000?101B1 000110000? 0011111100 000?0 0111011021 0000010010 0001100000 0111010001 001101A012 2000001111 0101020100 0011000200 1000?101D0 000101000? 0010?0???? 000?0 0100011120 00000000?0 1001100000 0111010001 0111011022 4000010001 1101030100 0111101200 1000?11121 000110000? 000101A101 000?0 0100111100 0000010011 0001100000 010??10000 ?00??0???? 0000100000 ??000????0 00?1000200 1001111110 100110000? 0000?1100? 000?0 0100111110 0000010011 1?00?????? ?????????? ??1101101? ??00000000 ??000????0 00?1000200 1001100111 000110000? 001111105? 000?? 0100111021 0000010010 1001100000 0111010001 0010010E22 0000110001 00010220?0 0111100000 1000?101D2 0001?10011 1000?11100 000?? 0100111100 0000010011 0000?00000 0011010001 0011011012 0000000001 0001032101 0001?00000 1000?00122 000110000? 0010?0???? 000?5 0100011021 0000010011 0001100000 010??10001 001101102? 0000000001 00010230?0 0000000201 1001101120 0001?1000? 0210?0???? 000?H 0100001021 0000000010 0001100000 000??00000 ?010211000 0000010000 ??01020100 0110000200 1000?00110 001101000? 0010?0???? 0011? 0100111110 0000010010 1??0?00000 000??10000 ?01001101? 0000000110 ??00120100 0110000100 1000?1111? 0001100011 0011110??? 000?? Taxon: zebralis gordialis Character rufdiscalis favibasalis primordialis Desmia tages Pyralis farinalis Pyralis Arnia nervosalis Midila guianensis Eporidia dariusalis Synaphe punctal is Conchylodes zebra Conchylodes Agrotera Botyodes diniasalis Ategumia ebulealis Cadarena pudoraria Cadarena Sufetula diminutalis Sufetula Aristebulea principis Arthromastix lauralis Arthromastix Bocchoris inspersalis Eudonia truncicolella Asciodes cf. Agathodes designalis Agrioglypta excelsalis Azochis cf. Crambus uliginosellus Crambus Dichocrocis cf.Dichocrocis Aetholix cf. Arthromastix pactolalis Arthromastix Asturodes fmbriauralis Asturodes Antigastra Eurrhyparodes lygdamis Eurrhyparodes Conogethes pandamalis Schoenobius gigantellus Cnaphalocrocis Diasemiopsis leodocusalis Anageshna cf. Apilocrocis novateutonialis Apilocrocis Multistate character states: A – (0&1); B (0&2); C (0&3); D (1&2); E (1&3); F (1&5); G (2&3); H (1&2&3). A 2. Multistate character states: Table

154 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019 0100111021 0000010010 ?001100000 000??10001 0010010002 4000001101 0000132101 0001000200 1000?10122 0001100010 0011011100 000?? 0100111101 000001000? ???0?00000 0111010000 ?011011000 0000000000 ??010320?0 0010000000 1000?001B? 0001?0000? 0010?0???? 000?? 0000110020 00001?0000 0000?00000 100??01010 ?0112????? ??00?00000 ??000????0 00?0000010 1000?00120 0011?0000? 000100???? 000?? 0100111101 0000010010 0001100000 010??10001 0011110010 1000120001 0000122100 0010000200 1000?10100 1001000110 001101101? 000?0 0100111100 00000100?0 0001100000 0011000000 ?01001032? 0000000001 1001030100 0010000200 1000?101D2 0001?1000? 000100???? 000?? 0100001120 0000010010 1001100000 0111010001 001101A222 0000010101 00010220?0 0111100200 1000?10100 000110000? 0000?11100 100?? 0100111020 00?0010010 1001100000 0111010001 001101AE02 2100020011 00010220?0 0110100200 1000?101B0 000110000? 0010?0???? 000?? 0100001120 0000010010 0001100001 0111010001 0011011012 2000010001 11000????0 01?0100200 1000?101D1 0011?0000? 0011011100 000?C 0100011001 0000010010 1000?00000 000??00001 0010010012 0100001111 0000122100 0011010200 1001000010 000110000? 0010?11111 000?0 0100001021 0000010010 1000?00000 0111010001 0010010012 4100001101 0100120100 0000000100 1000?00121 000100000? 0000?1101? 000?? 010011102? 0000010010 0001110000 010??10001 0010010002 4100020101 1100122100 0010000200 1000?001D0 0001100011 1000?1100? 000?? 0100111020 00000?0?10 00011????? ?????????? ?????????? ?????????? ?????????? ?????????0 0001100010 1000?0???? 000?? 01001?11?0 00000100?0 0000?00000 0011010001 001101A2?2 0000000001 0000120100 0111000200 1000?0010? ?????????? ?????????? ????? 0101111021 0000010010 0001100000 0111010001 0011011322 2100?0?101 1100102100 0111100200 1000?101B0 000110000? 001101102? 000?? 01????11?0 00??01??11 10011????? ?????????? ?????????? ?????????? ?????????? ?????????0 000110000? 0000?1105? 000?3 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111 0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345 0100001020 1001010010 1001100000 0111010001 001000??02 0000000001 1101020000 0111100201 1000?10112 0001100011 0000?0???? 0010? 010???11?1 ????0???10 00011????? ?????????? ?????????? ?????????? ?????????? ?????????2 100110000? 0000?0???? 000?? 0100011120 0001010010 0001100000 0111010001 0?11010222 0000?00001 1100122100 0111101200 1000?101B1 0001000010 0000?10??? 000?0 0100111120 0000010010 1001100000 0111010001 0011010222 4?00020001 1100120100 0111100200 1000?10100 0001100011 1000?10??? 000?? 0100111111 00000100?0 0000?00000 000??00100 ?010011012 0000000000 ??000????0 00?0000000 1000?000?1 000110000? 0000?1103? 0010? 0100001020 0000010010 0001100000 010??00000 ?010010100 0000001101 0100122100 1001010200 1000?10121 000110000? 0001011100 000?0 0100111120 0000010010 0000?00000 000??10001 0010010012 0000110001 0000120100 0000000200 10010101D0 000110000? 0010?0???? 000?0 0100111121 000001001? 0??1000000 010??10000 ?010011010 0000110001 0000120100 0010000100 1000?101D1 000110000? 0001010??? 000?? 0100001121 0000010010 0001100001 0111010001 0011011021 0100010011 100012G101 0110100100 1000?01120 001110000? 0000?0???? 000?? 0100011011 000001001? 0??0?00000 000??10000 ?00??????? ??01200000 ??00122000 0110010000 10011001B0 001100000? 0011011143 00101 0110011100 0000010010 0001100000 000??10001 0011011012 0000200001 11000????0 00?0100100 1000?11100 000110001? 000111105? 000?? 01000?10?0 00000100?1 0??1100001 0111010101 1011011001 1000000001 1110103000 0120000B00 1000?001D? ?????????? ?????????? ????? 0100011111 0000010000 0000?00000 000??10000 ?011011010 2000010010 ??010100?0 0100000000 1001001121 0001?1000? 0000?0???? 000?3 0100011020 0000010010 0000?00000 0111010001 0011011012 0000100001 1101022100 0110100200 0000?10121 000110000? 000100???? 000?? 0100001121 0000000000 0??1100000 000??00000 ?010011002 0000010000 ??001200?0 0010000000 1000?01120 000110000? 0000?0???? 000?A 0100011100 00000000?0 0001100000 0010010000 ?011010300 0000200000 ??001020?0 0110000100 1001101111 001110000? 0000?0???? 00113 0100011120 0000011010 1001100000 01110??001 0011211010 ??00010001 10000????0 01?1000000 1000?101D0 0001000010 001101100? 000?0 0100011100 0000010010 00011????? ?????????? ??1121101? ??00010101 00000????0 00?0000100 1000?00112 0001000010 001101100? 000?? 0100011101 0000010010 1000?00000 0111010001 001101A102 0000?01101 11010220?0 0110100001 1000?11122 000110000? 000101A111 0011G 0100111120 0000010010 0001100000 000??10001 0010011012 0000000001 00001200?0 0011000200 1001010110 0001110010 0000?1101? 000?0 0100111100 0000010010 0001100000 000??00000 ?01001A012 0000010010 ??001220?0 0000000200 1000?000?0 000110000? 0000?10??? 000?? 0100011100 0000000000 0001100000 0011010001 001121101? 0000010111 0000122100 0001000100 1001010122 001110000? 001100???? 000?? 0100011020 0000010011 0000?00000 0010010001 0011010022 2100?00001 0000120101 0110101200 1000?10120 000110000? 001101100? 000?? Taxon: silicalis pusialis zephyralis Character rubrocinctalis Patania ruralis Patania androstigmata Palpita Phostria temira Mecyna lutealis Mecyna Hydriris ornatalis Prenesta scyllalis Prenesta Lineodes vulnifca Liopasia andrealis Prenesta iphiclalis Prenesta Patania cf.Patania pseudocontortalis Hileithia obliqualis Nacoleia insolitalis Marasmia poeyalis Marasmia Hodebertia testalis Glyphodes sibillalis Omiodes humeralis Filodes fulvidorsalis Marasmia trapezalis Marasmia Leucochroma corope Leucochroma Metasia suppandalis Obtusipalpis pardalis Megastes cf. Glyphodes cf . stolali s Leucinodes africensis Omiodes continuatalis Niphopyralis chionesis Niphopyralis Lamprosema cf. dorisalis cf. Lamprosema Gonocausta cf. Neoleucinodes dissolvens Ghesquierellana hirtusalis Ghesquierellana Prenesta cf.Prenesta Prophantis cf.Prophantis Herpetogramma phaeopteralis Herpetogramma Neoanalthes cf. Table 2 continued. Table

155 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111 0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345 01000?11?0 00000100?0 0001100000 0111010000 ?110011011 0000?00001 0001032101 0000100200 1000?101D? ?????????? ?????????? ????? 0100011001 0000000010 0001100000 0111010000 ?010210010 0000000100 ???1031100 0110000000 1001010122 000000000? 001101103? 000?? 0100010111 00?0000010 0001000000 0110010000 ?011011000 0000000010 ???0120100 0010000000 10010001B0 000110000? 0000?1103? 000?? 0100011021 0000010010 1001100002 0111000001 0010010312 20000?1111 0101020100 0111000200 1001010100 000110000? 000100???? 000?? 01000?11?0 0000010??? 1001000000 0111010000 ?011211012 0000?31101 11000????0 10?1000200 0000?111D? 0001000010 001101100? 000?0 0100111121 0000010010 00010????? ?????????? ?????????? ?????????? ?????????? ?????????2 100110011? 1010?0???? 000?? 0100111121 0000010010 0001100100 0011010001 0011111010 0000020001 0000120100 0011101201 1001010110 000100000? 000111101? 000?? 0000000020 0000000010 0000?00000 000??00000 ?011211010 2100000010 ??00120100 0000000200 1000?10121 011110000? 00010110?? 000?? 0100001021 0000011010 0001100000 010??00001 0011111321 0000000101 00010030?0 0001000100 1001010122 0000?0000? 0010?11100 000?? 0110001000 0000010010 0001100000 0111010001 0010011102 2100100001 1100120120 0000100210 1000?101D2 000010000? 0010?1A05? 000?0 0100111111 0000010010 0000?00000 010??00100 ?01001101? 0000000000 ??000????0 00?0000000 10010001B2 000010000? 000101102? 0010? 0100001020 0000010010 1001100000 0111010001 00102110?? ??00001001 0000122100 0111000200 1000?1112? 0011100011 0000?1105? 000?? 0100111120 0100010010 1001101000 0111010001 001000??1? 4100100001 0001020101 0111100100 0000?10120 100110000? 000100???? 000?G 0100101021 0000010010 1001100000 011001000? ?011011012 2100100001 0000122101 0010000200 0000?11120 000110000? 0010?1102? 000?? 0100011100 0000010010 1??0?00000 010??00000 ?01101100? 0000100000 ??00120100 0010000000 10010001B2 0001000011 0000?1102? 000?0 0100011100 0000000001 1000?00000 000??10000 ?011011002 0000100010 ??00120100 0010000000 1000?01122 0001010011 100111102? 000?? 01000?11?0 00000100?0 1??1000000 000??10000 ?01101100? ??00110010 ??00120100 0020000000 1000?0011? ?????????? ?????????? ????? 01000111?1 0100010010 0001000000 010??10000 ?011011020 0000?00001 0001020100 0111000201 1000?00112 0001000011 0000?0???? 000?0 1100011100 0000010111 01?1100000 0?10010000 ??10010002 3000200001 0000110100 0110000100 1000?011D0 0001000011 0000?11130 00100 0100001101 0010010110 11?1100000 0111010000 ?010010102 0010010000 ??00131110 0110000100 10011011D2 0011001010 ?00111103? 00100 0100011120 0010010110 11?1100000 0111010000 ?010010102 0000010000 ??00110110 0110000100 10010011D2 000100100? 0000?1103? 00100 0100111120 0000010110 11?1100000 2111010000 ?01100??02 3000200001 1001023100 0010000100 11011011D0 0001010011 100111103? 010?? 0100?11101 00?0000110 01?10????? ?????????? ??110?2?12 2100?00001 00000????0 00?0?00200 10011?01?0 0001000011 000111A06? 010?? 0100111121 0000000110 0??0?00000 0111010001 0011012012 ??00100001 00000????0 00?0000200 00010001D? 0101100011 1000?1106? 010?? 0100111101 0000010111 11?1100000 0111010000 ??1110??0? 4100?00001 0000113100 000000??00 1101100120 0001?0000? 0000?1103? 010?? 01001?11?0 00000101?1 01?1100000 0111010000 ?011011002 0000110001 0001010110 0000000100 1000?001D0 000110000? 000111103? 0010? 1100011121 0000010110 01?1000000 0111010000 ?010011012 1?00010010 ??00101100 0010000100 10010011D0 0011000010 000111103? 00100 01001?11?0 00100101?1 01?1100000 0?0??10000 ?010010102 0000210001 0000131110 0110000100 1000?001D? ?????????? ?????????? ????? 0100011100 0000010111 11?1100000 0011010000 ?010110002 3100000001 00001G0110 0111000100 00010001B2 0001?1001? 0000?1103? 0010D 0100001100 0000010111 11?1000000 0?11010000 ?01001100? 0010110001 0000120110 0020000000 10011001D0 100110000? 000111102? 0010? 0100001100 00?0010011 01?1100000 0011010000 ?010010002 0000210011 0000120100 0011000100 10011001D? 000110000? 000111103? 00100 0100111121 0000010011 11?1100000 0111010000 ?11100??20 3000100001 10001230?0 0000000000 1100?001D2 0001000011 001101102? 0011? 0100111101 0000010111 01?1000000 0111010000 ?010010102 0000110000 ??00100110 0011000100 10011011D2 0011100011 0000?1103? 0010D 0100001120 0000010111 01?1100000 0011010000 ?010010102 0000110000 ??00111110 0100000100 10010001 Taxon: rantalis minima Siga liris astrigalis Character taeniolalis multiplicalis Oenobotys sp. Pagyda salvalis Pagyda Syllepis marialis Ostrinia nubilalis haemorrhoidalis Syngamia forella Zebronia phenice Zebronia Uresiphita Tetridia vinacealis Tetridia Trichaea pilicornis Trichaea Achyra cf.Achyra Anania verbascalis Euclasta gigantalis Sitochroa Cryptosara caritalis Cryptosara Pyrausta Spilomela perspicata Euclasta splendidalis Udeoides muscosalis Hyalobathra illectalis Hyalobathra Udea washingtonalis Paracorsia repandalis Paracorsia Terastia meticulosalis Terastia Loxostege aeruginalis Samea cf. Pycnarmon pantherata Hyalorista cf.Hyalorista Rhimphalea cf. Rhectosemia multifarialis Prophantis xanthomeralis Prophantis Salbia cf. Portentomorpha xanthialis Portentomorpha Pseudopyrausta cf.Pseudopyrausta Table 2 continued. Table

156 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

46 Presence of chaetae on surface of uncus head(s): (0) 63 Presence of a pair of sclerotized, hair-studded hair- absent (Fig. 15C); (1) present (Fig. 3D). pencils articulating with the anteromedian edge of 47 Structure of uncus head chaetae: (0) simple, not api- the saccus: (0) absent (Fig. 5E); (1) present (Fig. cally split (Figs. 7D, 14C); (1) bifurcate (Fig. 9I); 3G). (2) multifurcate (Popescu-Gorj & Constantinescu 64 Presence of fbula emerging from central inner val- 1977: fg. 7d); (3) short, fat, spatulate (Fig. 14C). va: (0) absent (Fig. 3D); (1) present (Fig. 4F). 48 Location of setae on uncus: (0) dorsal (Fig. 13A); 65 Presence of fbula emerging from dorsal valva base (1) dorsal & lateral (Fig. 5E); (2) dorsal & ventral near costa base: (0) absent (Fig. 3D); (1) present (Mally & Nuss 2010: fg. 2B); (3) ventral (Fig. 9E); (Figs. 4E, 10C). (4) lateral (Slamka 2008: pl. 29 fg. 182). 66 General shape of fbula: (0) broad triangular (Fig. 49 Attachment of uncus to tegumen: (0) broad, point of 8B); (1) elongate triangular, at least twice as long attachment constricted (Fig. 4E); (1) broad, smooth as broad (Fig. 4F); (2) elongate needle-like to claw- transition (Fig. 4F); (2) narrow, offset (Munroe shaped (Figs. 4E, 7D); (3) as long as broad, circular 1976b: pl. u fg. 6a; Mally & Nuss 2010: fg. 2C). to squarish (Fig. 12C). 50 Region between subscaphium and dorsal tegumen: 67 General orientation of fbula: (0) ventrally directed (0) membranous (Fig. 9G); (1) sclerotized as gnathos towards sacculus or distal sacculus (Figs. 4E, 8B); (sensu Maes 1998) (Fig. 3G); (2) sclerotized as pseu- (1) directed towards ventral sacculus base (Fig. 7C); dognathos (sensu Maes 1998) (Figs. 4E, 5D). (2) directed towards distal valva (Fig. 3A); (3) di- 51 Shape of transtillum arms: (0) triangular (tapering rected dorsally, towards tegumen/uncus (Fig. 3G). towards apex) (Fig. 4E); (1) rounded (Fig. 3G); (2) 68 Presence of chaetae on fbula surface: (0) absent strap-like (apex blunt or pointed) (Figs. 8B, 11E); (Fig. 4E); (1) present (Figs. 7C, 15D). (3) large rectangular, medioventrally with fnger-like 69 Structure of apex of chaetae on fbula surface: (0) simple (Figs. 7C, 15D); (1) some simple, some mul- process (“transtillum inferior” sensu Marion 1954) tifd (= editum of Pyraustinae) (Yang et al. 2012: up- (Fig. 15D; Marion 1954: fg. 11); (4) rhomboidal (= permost arrow in fg. 7A – D); (2) spatulate (Clarke triangular with cut apex) (Fig. 15C). 1986: fg. 34a). 52 Connection point of transtillum arms: (0) narrow 70 Presence of raised ridge running from basal to dorso- (Fig. 3G); (1) broad (Fig. 11E). distal sacculus: (0) absent (Fig. 5D); (1) present (Fig. 53 Presence of long dorsad chaetae on surface of tran- 12D). stillum arms: (0) absent (Fig. 3G); (1) present (Fig. 71 Presence of fnger-like process studded with simple 15D). chaetae emerging from central sacculus: (0) absent 54 Presence of lobar processes carrying hair-pencils (Fig. 5D); (1) present (Fig. 9H). on the dorsolateral tegumen sides: (0) absent (Fig. 72 Presence of extension (process in some cases) at dor- 13B); (1) present (Fig. 3A). sodistal sacculus: (0) absent (Fig. 4E); (1) present 55 Depth of gap/split of juxta: (0) < 10% of dorsoven- (Fig. 4F). tral length of juxta (Fig. 3G); (1) 10 – 60% of dorso­ 73 Spatial association of fbula with dorsodistal saccu- ventral length of juxta (Fig. 5D); (2) > 60% of dorso­­ lus (or its extension): (0) distant (Fig. 4F); (1) closely ventral length of juxta to complete division into two associated, overlapping (Fig. 4E); (2) fbula and dor- juxta arms (Figs. 3A, 6E). sodistal sacculus fused (Fig. 11E). 56 Saccus shape: (0) U-shaped (Fig. 3A); (1) (sharply) 74 Presence of infation of basal costa: (0) absent (Fig. V-shaped (Fig. 4F); (2) stout, almost rectangular 4E); (1) present (Fig. 8B). (Fig. 5E); (3) narrow elongate (Fig. 9H). 75 Joint of basal valva costa (with vinculum) extended 57 Presence of constriction at basal saccus: (0) absent into an elongate, rod-shaped process: (0) absent (Fig. (Fig. 3A); (1) present (Fig. 13A). 4F); (1) present (Fig. 8B). 58 Ratio between saccus length and sacculus breadth: 76 Presence of long, sometimes loosely arranged chae- (0) ≤ 1 (Fig. 3A); (1) > 1 (Fig. 13B). tae on surface of costal base: (0) absent (Fig. 4E); (1) 59 Presence of protruding keel on ventral saccus tip: (0) present (Figs. 3A, 5E). absent (Fig. 3A); (1) present (Fig. 4F). 77 Presence of a knee-like bend of 60 – 80° in the post- 60 Presence of partly sclerotized, chaetose hairpencil basal costa: (0) absent (Fig. 4E); (1) present (Fig. articulating with the anterior edge of the vinculum- 13B). tegumen connection: (0) absent (Fig. 7C); (1) pre- 78 General shape of post-basal costa (not the entire dor- sent (Figs. 13A, 10C, 14E). sal valva edge): (0) concave (Fig. 3A); (1) straight 61 Number of hairpencil sclerites on each side of the (Fig. 5E); (2) convex (Figs. 8B, 9F,G). genitalia: (0) one (Figs. 13A, 10C, 14E); (1) two or 79 Presence of a setose dorsad process on the basal cos- more (articulated with each other via membranes) ta: (0) absent (Fig. 3A); (1) present (Fig. 11E). (Fig. 9H). 80 General structure of distal costa: (0) tubular (Fig. 62 Presence of more than one kind of hairpencil chae- 3A); (1) broadening (Fig. 12E). tae: (0) absent (Fig. 14E); (1) present (Clarke 1986: 81 Costa following the course of (= in alignment with) fg. 34a; Kimura et al. 2002: fgs. 1 – 4). the dorsal valva edge (all the way) into subapical

157 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

valva region: (0) absent (Fig. 9H); (1) present (Fig. 101 Presence of thickened mesocuticle in the collicu- 4E). lum: (0) absent (Fig. 5F); (1) present (Fig. 14G). 82 Presence of detached costa from valval area, the 102 Point of attachment of ductus seminalis to female costa protruding freely dorsad instead: (0) absent genital tract: (0) at posterior ductus bursae, at or (Fig. 7C); (1) present (Fig. 15C; Munroe 1976b: near colliculum (Fig. 5G); (1) at anterior ductus pl. u fg. 6a; Shaffer & Munroe 2007: fgs. 130, bursae (Fig. 7E); (2) at corpus bursae (Fig. 12F). 133). 103 Demarcation between corpus bursae and ductus 83 Presence of a recess in the course of the ventral val- bursae: (0) distinct by narrow anterior ductus trans- va edge: (0) absent (Fig. 4F); (1) present (Fig. 7D). forming into wide corpus bursae (Fig. 3E); (1) 84 Presence of a coecum on the phallus apodeme: (0) indistinct or absent by wide anterior ductus trans- absent (Figs. 8C, 12D); (1) present (Figs. 3B, 4G). forming into equally wide corpus bursae, i.e. fuent 85 Length of phallus coecum relative to phallus apo­ transformation of d.b. to c.b.) (Figs. 3F, 10E). deme length: (0) < 10% (Fig. 4G); (1) ≥ 10% (Fig. 104 Presence of sclerotisation in ductus bursae: (0) ab- 3B). sent (Fig. 3E); (1) present (Fig. 5F). 86 Presence of reduction of phallus apodeme scleroti- 105 Intensity of ductus bursae sclerotisation: (0) weak sation to a ventral, longitudinally sclerotized strip (granulose texture) (Fig. 13E); (1) strong (Fig. 5F). (the rest of the apodeme being more or less membra- 106 Presence of sclerotisation in corpus bursae: (0) ab- nous): (0) absent (Figs. 4G, 15D); (1) present (Figs. sent (Fig. 9J); (1) present (Fig. 3F). 7D, 10C, 13F). 107 Structure of corpus bursae sclerotisation: (0) a gran- 87 Presence of a distinct sclerite in the posterior phal- ulose area (Figs. 11G – H); (1) one or more clearly lus apodeme: (0) absent (Fig. 13F); (1) present (Fig. delimited sclerites (= signum, Pl. signa) (Fig. 3F). 4G). 108 Number of signa: (0) one (Fig. 3E); (1) two or more 88 Presence of sclerotisation on surface of vesica: (0) (Fig. 3F). absent (Fig. 7F); (1) present (Fig. 3B). 109 Shape of anterior-most signum: (0) circular, 89 Type of vesica sclerotisation: (0) single cornutus spinose, can be invaginated as a spine (Fig. 8K); (1) (Fig. 3B); (1) multiple cornuti (Fig. 13F); (2) granu- longitudinal slim, strip-like (Fig. 13E); (2) elongate lated area (Fig. 3B). rhombical to ovate (longitudinal axis longer than 90 Orientation of everted papillae anales: (0) postero­ transverse one) (Fig. 5F); (3) transverse rhombical ventrad (Fig. 3F); (1) ventrad (Fig. 4H); (2) poste- to cross-shaped (longitudinal axis shorter than or riad (Fig. 5F). equally long as transverse one) (Figs. 3E, 15F – G); 91 Dorsal end of papillae anales larger than ventral (4) patch of protruding teeth/spikes (Fig. 3F); (5) end: (0) absent (Fig. 3F); (1) present (Fig. 13E). transverse, smooth or dentate line or arch, with or 92 Ventral end of papillae anales larger than dorsal end: without central posteriad leg (if present, then sig- (0) absent (Fig. 3F); (1) present (Fig. 14G). num Y-shaped) (Figs. 10E, 11H); (6) broad, medi- 93 Presence of a strongly sclerotized frame (= lamella ally constricted, resembling puckered lips (Figs. antevaginalis) around the ostium bursae: (0) absent 14G,H). (Fig. 5F); (1) present (Fig. 3F). 110 Shape of second signum (located posterior of frst 94 Presence of strong sclerotisation in the antrum: (0) signum): (0) circular, spinose, can be invaginated absent, with antrum more or less membraneous as a spine (Fig. 8K); (1) longitudinal slim, strip-like (Fig. 11H); (1) present (Fig. 4H). (Fig. 5F); (2) elongate rhombical to ovate (Shaffer 95 Presence of a longitudinal membranous strip in the & Munroe 2007: fgs. 299, 300); (3) patch of pro- antrum sclerotisation: (0) absent (Fig. 9J); (1) pre- truding teeth/spikes (Fig. 3F). sent (Figs. 5F, 10D,E). 111 Presence of a third, slim, strip-like signum posterior 96 Presence of thickened mesocuticle in the antrum: of the two anterior signa: (0) absent (Fig. 8J); (1) (0) absent (Fig. 11G); (1) present (Fig. 4H). present (Fig. 8K). 97 Presence of a cone-shaped central structure (Ana­ 112 Presence of appendix bursae on anterior ductus bur- nia-type) in the antrum: (0) absent (Fig. 11G); (1) sae: (0) absent (Fig. 8K); (1) present (Figs. 14G, present (Fig. 15F; Tränkner et al. 2009: arrows in 15G). fgs. 18 – 21). 113 Presence of appendix bursae on corpus bursae: (0) 98 Presence of a lateral blind-end evagination (diver- absent (Fig. 8K); (1) present (Figs. 3F, 15F). ticulum) in the colliculum: (0) absent (Fig. 13D); (1) 114 Point of attachment of appendix bursae on corpus present (Fig. 13E). bursae: (0) lateral (Figs. 3F, 15F); (1) posterior (Fig. 99 Presence of a strongly sclerotised colliculum ante- 9J); (2) anterior (Fig. 5G). rior of antrum and posterior of attachment of ductus 115 Locality of larval feeding: (0) concealed in rolled/ seminalis: (0) absent (Fig. 11G); (1) present (Fig. spun or in a web (Leuthardt et al. 2010: fg. 5F). 1; Hayden et al. 2017: fg. 19); (1) on / sur- 100 Presence of a longitudinal membranous strip in the face (upper/underside) (Hayden et al. 2013: Line­ colliculum sclerotisation: (0) absent (Fig. 13D); (1) odes fontella); (2) boring in stems and/or branches present (Fig. 5F). (Sourakov 2011: fgs. 6A, 7A, 10B); (3) boring in

158 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

fowers, pods and (Sourakov 2011: fgs. 3, nae) is sister to the “CAMMSS Clade” sensu Regier et 4); (4) on/in roots (Genty & Mariau 1975: fgs. al. (2012), with a clade Crambus (Crambinae) + Eudonia 3 – 5, 7); (5) on dead and decaying plant matter (Scopariinae) sister to the “Wet Habitat Clade” (sensu (Murphy 1990: pl. 15 fg. J). Regier et al. 2012) Clepsicosma + (Midila + Schoeno­ bius), the latter belonging to Acentropinae, Midilinae and Schoenobiinae, respectively. Sister to the Crambidae 3.3. Phylogenetic results outgroup is the “PS Clade” (sensu Regier et al. 2012) of Pyraustinae and Spilomelinae. Both Pyraustinae and Spilomelinae are highly supported (1 PP) monophyletic The investigation of the gene data with DAMBE5 showed and moderately-supported (0.93 – 0.95 PP) sister to each no signs for signifcant substitution saturation in the other. three different codon positions of COI, CAD, EF-1a and Within Pyraustinae, Tetridia is sister to all other taxa. GAPDH. Codon positions nt1 and nt2 of IDH and RpS5 The two Euclasta species form a monophylum (Euclas- showed no signifcant saturation, whereas in nt3 of these tini) that is sister to the remainder of Pyraustinae. A clade two genes, some substitution saturation was observed. Uresiphita + (Portentomorpha + (Cryptosara + Hyalo­ This low level of substitution saturation was accepted as bathra)) is sister to the remainder of Pyraustinae (Pyraus- of minor infuence for the phylogenetic analysis of these tini). Ostrinia is sister to a clade Pagyda + Paracorsia, data, so that no codon position was removed from the which is sister to the remainder of Pyraustinae. Achyra + fnal dataset. This decision was supported by trial analy- (Loxostege + Sitochroa) is sister to a clade Oenobotys + ses of the dataset with nt3 removed from IDH and RpS5 (Hyalorista + Pyrausta) and its sister group of Psammo­ which showed a very similar topology and comparable tis, Pseudopyrausta and Anania. Anania, with two sam- node support. pled species, is monophyletic except in the phylogram of In phylogenetic pre-analyses, several taxa had con- the GENES-partitioned genetic dataset, where A. hortu­ spicuously unstable positions in the phylogeny. These lata is sister to Psammotis, and A. verbascalis sister to most problematic ‘rogue’ taxa were identifed using Pseudopyrausta, all with PP < 0.9. RogueNaRok and excluded from fnal analyses. One ex- The phylogenetic relationships within Spilomeli- ception was Niphopyralis chionesis, which we decided to nae are as follows: Hydririni + ((Udeini + Lineodini) + keep in the dataset despite its long terminal branch in the (Wurthiini + (Agroterini + (Margaroniini + (Spilomelini phylogenetic results. + (Herpetogrammatini + ((Hymeniini + Asciodini) + Differences in the coverage of morphological data (Trichaeini + (Steniini + Nomophilini))))))))), with the coding affect the performance of the phylogenetic analy- exception of the GENES-partitioned analyses, where ses: MrBayes analyses containing morphological data Spilomelini is in an unsupported (0.67 – 0.72 PP) sister- for the outgroup taxa performed worse than analyses group relationship with Margaroniini (indicated by dotted that only comprised morphological data for Pyraustinae arrow in Fig. 1). Hydririni comprises Hydriris + (Lam­ and Spilomelinae and where outgroups were coded as ‘?’. When outgroup morphological data is included in prosema + (Gonocausta + Syllepis)). Udeini comprises the analysis, the parallel MrBayes runs do not converge Conchylodes + (Udeoides + Udea). Lineodini comprises properly and the effective sample size is low for a num- Lineodes + (Rhectosemia + (Leucinodes + Neoleuci­ ber of parameters. All phylogenetic results stated and nodes)). Wurthiini comprises Apilocrocis + (Aristebu­ discussed below are therefore based on the datasets that lea + Niphopyralis). Agroterini comprises Pycnarmon + only comprise the morphological data for Spilomelinae, ((Neoanalthes + (Aetholix + Agrotera)) + (Haritalodes Pyraustinae and Sufetula. The potential causes and impli- + (Phostria + Patania))). Margaroniini forms a large cations of outgroup coding are elaborated in the Discus- polytomy with several moderately to well-supported sion section. monophyla, which are: Asturodes + Maruca; Omiodes; The parallel runs of all MrBayes analyses converged Prenesta; Liopasia + (Agathodes + Terastia); Hodeber­ suffciently after 30 Mio. generations, and ESS were tia + (Antigastra + Zebronia); (Azochis + Conogethes) + (mostly well) above 100. The analyses of the different Ghesquierellana + Megastes); (Agrioglypta + Obtusipal­ datasets result in highly similar topologies, and branch pis) + (‘Dichocrocis’ cf. zebralis + Glyphodes). Addition- support from the analyses of the molecular data alone al taxa in Margaroniini with unresolved or unsupported and those of the combined molecular and morphological (PP < 0.9) relationships are: Cydalima, Filodes, Rhim­ data are almost identical (Fig. 1). The additional mor- phalea, Diaphania, Palpita, Botyodes, Cadarena, Leu­ phological data in the analysis of the combined dataset cochroma, ‘Nacoleia’ insolitalis. Spilomelini comprises does not result in improved resolution or branch sup- (Siga + Eporidia) + (Spilomela + (Salbia + (Marasmia + port as compared to the results of the molecular data- Cnaphalocrocis))). Herpetogrammatini comprises Eur­ set. Branching differences are found in the position of rhyparodes + (Herpetogramma + Hileithia). Hymeniini the clade Spilomelini (see dotted arrow in Fig. 1), and comprises Hymenia + Spoladea. Asciodini comprises within the clade Margaroniini. Pyralidae, Pyralinae, and Asciodes + Arthromastix. Trichaeini comprises Trichaea Crambidae are each monophyletic with high branch sup- + Prophantis. Steniini comprises (Dolicharthria + Meta­ ports. In the Crambidae outgroup, Sufetula (Lathroteli- sia) + (Duponchelia + Anageshna). Nomophilini com-

159 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 1.

prises (Syngamia + (Ategumia + (Bocchoris + (Diasemia The topology of the outgroup CAMMSS clade is differ- + Diasemiopsis)))) + (Desmia + (Mecyna + (Samea + ent than that in Regier et al. (2012). Euclastini diverges Nomophila))). frst in Pyraustinae. The second-diverging clade is Por- Parsimony analysis resulted in three cladograms of tentomorphini including Portentomorpha, Hyalobathra, 35,100 steps. The strict consensus (Fig. 2) has 35,153 and Cry­pto­sara. The topology within Pyraustini is sub- steps. Condensing these cladograms with “collapse[” did stantially different. Tetridia and Uresiphita are subordi- not lengthen them, so fltering with the command “best” nate in Pyraustini, sister to Pseudopyrausta and Ostrinia, was not necessary. Niphopyralis groups with Sufetula. respectively.

160 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Fig. 1. Bayesian consensus phylogram of the three parallel runs of the TIGER-partitioned MrBayes analysis of the molecular+morphology dataset (“mol+morph-TIGER”). Numbers at internal branches are PP ≥ 0.9, above branches “mol+morph-GENES | mol+morph-TIGER”, below branches “mol-GENES | mol-TIGER”; nodes without posterior probabilities indicate PP ≥ 0.9 in all four analyses. Scale bar repre- sents substitutions per site. Clade names in quotation marks correspond to those in Regier et al. (2012). — Abbreviations: n/a – node not present; PP – posterior probability. — Symbols: - PP < 0.9.

The three unweighted cladograms differ in only two Diasemia and Diasemiopsis) consistently being sister to clades: 1) whether Lineodes or Rhectosemia is the frst- the clade of the other three tribes (Steniini, Trichaeini, diverging genus of Lineodini, and 2) the topology (Steni­ ­ and Nomophilini s.str.). ini + (Nomophilini + Trichaeini)) versus (Nomophilini + IterPCR (Pol & Escapa 2009) did not suggest any (Steniini + Trichaeini), with Nomophilini in a reduced taxa or characters to recode. Implied weighting with k- sense including Syngamia but not the Ategumia clade. parameter values of 9 through 13 found cladograms (not Other differences are 1) the grouping of Desmia with shown) with Niphopyralis sister to Aristebulea principis Trichaeini and 2) the Ategumia clade (with Bocchoris, Munroe & Mutuura, 1968, but the topologies are oth-

161 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 2. Strict consensus of three parsimony cladograms of 35,100 steps, with morphological characters mapped with slow optimization. — Symbols: ● unique apomorphies; ○ homoplastic apomorphies. erwise similar to the equal-weights results. The lengths cally longer than the shortest actual cladograms, the extra range from 35,159 to 35,192 steps. steps did not occur along the particular clades that we are Among the genera with more than one sampled spe- interested in diagnosing. cies, Euclasta, Udea, Patania and Omiodes are mono- phyletic. Marasmia, Prenesta and Samea are paraphyl- etic, and Glyphodes and Dichocrocis are polyphyletic. 4. Phylogenetic classification The morphological data were mapped with WinClada on the Bayesian consensus (Fig. 1, synapomorphies not shown) and the parsimony consensus (Fig. 2) using slow In this section we focus on the taxonomic circumscrip- optimization (= delayed transformation, DELTRAN). tion of Spilomelinae and Pyraustinae and the clades The results are stated in the diagnoses of the clades in found therein. We state synapomorphies and / or charac- the taxonomy section. Although consensus trees are typi- ters derived from slow optimization in the “Synapomor-

162 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Fig. 2 – Continuation. phies” paragraphs. Representatives of all proposed tribes apomorphies for the respective tribe. A checklist of all are illustrated in Figs. 3 – 15. The morphological charac- Spilomelinae and Pyraustinae genera that are placed in ters indicated in the fgures do not necessarily represent tribes is given in the Appendix.

163 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

4.1. Spilomelinae + Pyraustinae (PS) Clade this signum type with the minor transverse axis reduced or absent as “ediacaroid” signum, after the Ediacaran biota from the late Proterozoic Eon, which show similar Synapomorphies. No unambiguous synapomorphies body shapes that likewise vary from nearly circular to were found, as most of the Crambidae outgroup taxa elongate. were left uncoded. Under slow optimization, the fol- The corpus bursae has an appendix bursae. An ap- lowing three characters are found: 36:1, U-shaped scle- pendix is present in most investigated Pyraustinae and rotisation of lateral and anterior edge of male sternite 8 in most Hydririni as well as in Conchylodes and Sisyrac­ present (also present in Schoenobiinae: Schoenobius); era (Udeini). This character is absent in all other investi- 65:1, presence of fbula emerging from dorsal valva base gated Spilomelinae except for several Margaroniini and near costa base; 106:1, sclerotisation in corpus bursae Eporidia (Spilomelini), where it might be a secondary present. development. Description. The uncus has bifd chaetae. These dis- Remarks. Immature stages of Pyraustinae and Spilo- tinctive chaetae are present in many Pyraustinae as well melinae have not been studied in a phylogenetic context, as in most Spilomelinae. Bifd chaetae are lost in some and characters consistently separating the two groups are spilomeline groups which have normal, hair-like mono- not known (Allyson 1981, 1984). flament chaetae on the uncus instead (e.g. Conchylodes genus group in Udeini, Agroterini, several Margaro- niini). 4.2. Spilomelinae Guenée, 1854 The costa of the valva is straight to concave. All investigated Pyraustinae as well as the non-euspilome­- Type genus: Spilomela Guenée, 1854 line clades (Hydririni, Udeini, Lineodini, Wurthiini) ex- = Sylleptinae Swinhoe, 1900 hibit male genitalia with a straight or concave costa (we only refer to the costa here, and not to the whole dor- Synapomorphies. 10:1, maxillary palpi minute to ob- sal valva edge). Most other Spilomelinae have a convex solete, cannot hypothetically come in contact with each costa. other; 23:0, fornix tympani projecting in ventral direc- The gnathos is reduced to a transverse strap, laterally tion (unique); 105:0, ductus bursae sclerotisation weak fused to the tegumen, and usually without a central pro- or with granulose texture. cess. This is the “pseudognathos” of Maes (1998a), which Description. The fornix tympani projects ventrad from Solis & Metz (2011) homologized with the gnathos: the the tympanic frame. The retinacular hook (frenulum hook structure is simply reduced and fused. A few Pyrausti- sensu Forbes 1926) is lost. Females have two frenular nae and Spilomelinae do have a central process like that bristles, while the number of female frenular bristles var- in most other Crambidae, such as Munroeodes Amsel, ies in Pyraustinae. 1957, Sarabotys Munroe, 1964, Phaedropsis Warren, Systematics. Spilomelinae includes a monophylum that 1890, Patania Moore, 1888, Syllepte amando (Cramer, we refer to as “euspilomeline clade” (Greek eu- good, 1779), Deuterophysa Warren, 1889 and Mimudea War- true), characterised by two morphological synapo- ren, 1892. Munroe (1964) considered this process to be morphies (see below). In contrast, the tribes Hydririni, primitive and indicative of relationship with Evergestini, Udeini, Lineodini and Wurthiini represent a paraphylum, but our results indicate that such processes are secondar- of which Wurthiini is sister to the euspilomeline clade. ily derived and homoplastic. Because of this paraphyly, we refrain from proposing a The phallus apodeme is evenly sclerotized. All inves- name for the group, and refer to them as the non-euspi- tigated Pyraustinae (except Uresiphita) and the non-eu- lomeline clades. spilomeline clades exhibit this character. In the euspilo- melinae clades, the sclerotisation of the phallus apodeme is usually reduced to a longitudinal ventral strip stretch- 4.2.1. Non-euspilomeline clades ing the length of the phallus; this character is reversed in several Spilomelinae in the euspilomelinae clades. The non-euspilomeline clades are characterised by ple- The signum is rhombiform. This distinctive signum siomorphies shared with Pyraustinae: the pleural mem- is a traditional character of Pyraustinae s.str. (Munroe branes of the male abdominal segment 8 lack a longitu- 1976a). It is a single sclerite with two axes, a major and dinal sclerotized strip; the valva costa is straight or con- minor one, and has short spines or granules. Apart from cave; the phallus apodeme is evenly sclerotized; and the Pyraustinae, this signum type is found in modifed forms signum is “ediacaroid”. Several taxa exhibit an appendix in the non-euspilomelinae clades (see below). Lamprose­ bursae. ma victoriae Dyar, 1923 has a very rhombiform signum, as do other Laprosema spp. and Gonocausta sabinalis Dyar, 1914. In other Hydririni and in Udeini, the minor 4.2.2. Hydririni + Lineodini transverse axis is nearly absent, and the whole is elongate and zipper-shaped (Syllepis, Udea, Conchylodes, Rhec­ Synapomorphies. 9:1, intersexual size difference of 3rd tosemia) to nearly circular (); we refer to labial palpomere, short in male.

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Fig. 3. Hydririni. A: male genitalia of Hydriris ornatalis. B: phallus of H. ornatalis. C: adult male of Lamprosema sp., a representative of the core-Lamprosema group. D: male genitalia of Syllepis marialis. E: anterior part of female genitalia of Gonocausta sp. F: female genitalia of H. ornatalis. G: male genitalia of L. cf. dorisalis. H: posterior abdomen (spread) of L. cf. dorisalis. — Scale bars: A, B, D – H – 500 µm; C – 5 mm.

165 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

4.2.3. Hydririni Minet, 1982 stat.rev. Hydriris ornatalis larvae feed on the leaf undersides of batatas (Convolvulaceae) and related , Type genus: Hydriris Meyrick, 1885 later instars skeletonize the leaves (Hinckley 1964). Remarks. The genitalia of Syllepis and Gonocausta are Synapomorphies.113:1, appendix bursae present on cor- highly similar, and a future revision might evaluate these pus bursae (see remarks below). Most Hydririni exhibit two genera as congeneric. Lamprosema contains numer- the unique apomorphy 38:1, central hair scale patch(es) ous misplaced Old-World species and needs revision. We on anterior edge of male sternite 8 present. In the par- verify the congenerity of the taxon used in our analyses simony trees, only 113:1 is an apomorphy of Hydririni. with Lamprosema lunulalis Hübner, 1823 from Suri- Description. The genitalia morphology is heterogene- name, the type species of the genus. ous: the valvas are slender to broad (Syllepis), and the An appendix bursae is also observed in Conchylodes uncus and gnathos are reduced (Choristostigma War- and Sisyracera (Udeini) as well as in Pyraustinae. Un- ren, 1892, Hydriris Meyrick, 1885, Hyperectis Meyrick, der a slightly different basal branching sequence, the ap- 1904) to well-developed. Choristostigma, Hydriris, Hy­ pendix bursae could be recovered as a synapomorphy of perectis, Nehydriris Munroe, 1974a and Rhectothyris Pyraustinae and Spilomelinae but lost in most Spilomeli- Warren, 1890 have a dorsolateral tegumen exhibiting nae. However, the best topology in this study indicates lobar processes with a feld of long, thin hair-pencils, separate origins. with a long phallus caecum, and with a single straight or Some of the characters by which Minet (1982) placed hooked cornutus. The hairs on the anterior edge of male Hydririni in Glaphyriinae are homoplastic. Spatulate sternite 8 are absent in Choristostigma, Ommatospila hind wing scales are paralleled with Glaphyriinae, and and Hydriris ornatalis but present in H. aonisalis. Some Solis & Adamski (1998) found that such scales are vari- taxa have an appendix bursae emerging laterally from the able even within Glaphyriinae. The spinose signa of H. corpus bursae as in Pyraustinae: Pyraustini (see below). ornatalis resemble the spinose sclerotizations of many The signum is ediacaroid, circular to elongate with the Neotropical glaphyriines, but H. aonisalis has a lenticu- minor transverse axis varying from broad (Lamprosema lar ediacaroid signum. Hübner, 1823, Gonocausta Lederer, 1863) to short, or forming circle(s) of radiating spines (Choristostigma, Hydriris, Nehydriris). Ommatospila has a circular edi- 4.2.4. Lineodini Amsel, 1956 stat.rev. acaroid signum and an opposing signum consisting of a feld of spines, like in Choristostigma and Hydriris. Type genus: Lineodes Guenée, 1854 Systematics. Minet (1982) established Hydririni in Gla­ phyriinae and included only Hydriris in this tribe. Mun- Synapomorphies. 104:0, sclerotisation in ductus bursae roe (1995) returned Hydriris to Spilomelinae. Beside absent. Slow optimization only: 8:1, direction of 3rd labi- Hydriris, Munroe (1995) placed in his Hydriris genus al palpomere porrect (paralleled in other early-diverging group also Choristostigma, Dyar, 1906 and Ne­ clades; not found in the parsimony trees); 95:1, longi- hydriris. tudinal membranous strip in the antrum sclerotisation According to our phylogenetic analysis, Hydriris (7 present. Fast optimization only: 19:0, female with only spp.), Gonocausta Lederer, 1863 (4 spp.), Lamprosema one frenular bristle (Hayden et al. 2013); 106:0, signum (72 spp.) and Syllepis Poey, 1832 (7 spp.) belong to Hy- absent (not with slow optimization due to position of dririni. Furthermore, based on morphological characters Rhectosemia). we place Choristostigma (10 spp.), Nehydriris (1 sp.), Description. The wings are moderately broad (Leuci­ Ommatospila Lederer, 1963 (3 spp.) and Rhectothyris (1 nodes) to narrow and almost pterophorid-like in Lineodes sp.) here. According to morphological characters, Gesh­ Guenée, 1854 and Atomopteryx Walsingham, 1891. The na does not belong to Hydririni, but to Spilomelini (see sacci tympani are ventrally open (Hayden et al. 2013). below). The valvae are very slender to relatively broad, trian- Hyperectis dioctias Meyrick, 1904, the type species gular or paddle-shaped, and the valva apex is rounded of Hyperectis, is depicted in Zimmerman (1958). From to somewhat acute; the costa is straight to concave; the there it is evident that this genus is misplaced in Pyraus- fbula is either slender and emerging from the costa base, tinae and that the genitalia are close to those of Hydriris shorter and emerging more from the centre of the valva, ornatalis (Duponchel, 1832) and H. aonisalis (Walker, or entirely absent in Euleucinodes Capps, 1948 and Pro­ 1859), and that the genus is not distinguishable from leucinodes Capps, 1948 (see Capps 1948). The sacculus Hydriris. We therefore synonymize Hyperectis Meyrick, is simple or (in Leucinodes Guenée, 1854) with a dis- 1904 syn.n. with Hydriris, and transfer the two species tal sacculus process in close association with the fbula. Hydriris dioctias Meyrick, 1904 comb.n., and Hydriris The posterior phallus is unmodifed or with sclerotized apicalis (Hampson, 1912) comb.n. appendages (in Leucinodes, see Mally et al. 2015). The Food plants. The known larval food plants are Sapin- posterior ductus bursae, colliculum and antrum in Leu­ daceae (Gonocausta, Lamprosema, Syllepis) and single cinodes and Neoleucinodes Capps, 1948 often have a cases of Fabaceae (Lamprosema), and thickened mesocuticle and partial sclerotisation (Hayden (both Syllepis) (Janzen & Hallwachs 2009). et al. 2013; Mally et al. 2015).

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Fig. 4. Lineodini. A: ventral view of wings on frenulum bristle of female . B: head of Le. orbonalis, male (left) and female (right) (modifed from Figs. 11 – 12 of Mally et al. 2015). C: adult male of Lineodes vulnifca. D: adult female of Le. Africensis. E: male genitalia of Li. vulnifca. F: male genitalia of Le. africensis. G: phallus of Le. africensis. H: posterior part of female genitalia of Le. pseudorbonalis. — Scale bars: C, D – 5 mm; E – H (same scale) – 500 µm.

Systematics. Lineodini was originally proposed for Li­ our phylogenetic analysis as well as to contain Euleuci­ neodes (38 spp.) and Atomopteryx (10 spp.) (= Steno­ nodes (1 sp.) and Proleucinodes (4 spp.), and to confrm ptycha Zeller, 1863) (Amsel 1956); it is expanded here Atomopteryx based on morphological characters. to contain Leucinodes (20 spp.), Neoleucinodes (9 spp.) With the exception of Leucinodes, all these genera and Rhectosemia Lederer, 1863 (12 spp.) according to were included by Munroe (1995) in his Udea genus

167 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

group, plus Lamprosema and Udea, which however be- (infated in Cheverella), the valva apical of the sacculus long to Hydririni (see above) and Udeini, respectively tapers towards a rounded apex. The female genitalia have (see below). an elongate signum that is rhombical, lanceolate or edi- The position of Rhectosemia in the phylogram (Fig. 1) acaroid in shape. In all Udea species groups sensu Mally diverging after Lineodes disagrees with morphology. Fe- & Nuss (2011) except the U. ferrugalis species group, an males of some examined species of Rhectosemia have accessory signum in the conjunction of ductus- and cor- two frenular bristles, and they have a signum that is usu- pus bursae is present. Conchylodes, and Sisyracera ally narrow and elongate. These are plesiomorphies in have a membranous appendix bursae, attached anteriorly contrast to the synapomorphies of the other genera: one in Conchylodes and Ercta and posteriorly in Sisyracera. bristle in both sexes and the loss of the signum. The antrum is strongly sclerotized, weakly in Cheverella. Food plants. Almost all known larval food plants are Systematics. Minet (1982) associated Udea (214 spp.) , the larvae either boring into the fruits (Leu­ with Pyraustinae, a decision followed by Leraut (1997, cinodes, Neoleucinodes) or feeding on leaves and fruit 2012). In contrast, our phylogenetic analysis supports surfaces (Atomopteryx, Lineodes) (Hayden et al. 2013). Udea as belonging to Spilomelinae, forming a monophy- Several species of this group are pests on solanaceous lum together with Udeoides (5 spp.) and Conchylodes crops, e.g. Neoleucinodes elegantalis on (Sola­ (21 spp.); furthermore, based on morphological charac- num lycopersicum), and Leucinodes spp. on eggplant ters, we place Cheverella (1 sp.), (1 sp.), Ercta (7 (Solanum melongena) (Hayden et al. 2013; Mally et al. spp.), Mnesictena Meyrick, 1884 (7 spp.), Sisyracera (3 2015). Janzen & Hallwachs (2009) report two Neoleuci­ spp.) and Tanaophysa (2 spp.) in this monophylum. nodes species from Heliconia spp. (Heliconiaceae). Udeini was proposed by Leraut (1997) in Pyrausti- Remarks. Character 95:1, the presence of a longitu- nae, but without a description to differentiate the taxon, dinal membranous strip in the antrum sclerotisation, a requirement by the International Code of Zoological is also present in Hydririni: Gonocausta, Syllepis. The Nomenclature (Ride et al. 1999, International Commis- reduction to one frenular bristle in females is also pre- sion on Zoological Nomenclature: article 13.1) for names sent in members of the Udea itysalis and U. alpinalis published after 1930. Therefore, the family-group name species groups (sensu Mally & Nuss 2011) (Udeini), in Udeini was not available prior to our proposal and formal Metasia suppandalis (Steniini), Diasemiopsis ramburi­ description. alis (Duponchel, 1833) (Diasemiini) and Niphopyralis The genitalia of Azochis graphialis Schaus, 1912, (Wurthiini). We fnd Lineodini and Udeini (see below) to type species of Nonazochis Amsel, 1956, resemble those be sister groups in the Bayesian analyses. However, they of C. diphteralis (Geyer, 1826), not justifying the sepa- do not share any synapomorphies with each other, also ration of the two genera. We therefore synonymize the not under slow optimization. Lineodini and Hydririni are monotypic Nonazochis Amsel, 1956 syn.n. with Conchy­ sister-groups in parsimony analysis, sharing 9:1, 68:0, lodes Guenée, 1854, and transfer Conchylodes graphialis 99:0, and 109:3. (Schaus, 1912) comb.n. Conchylodes octonalis (Zeller, 1873) comb.n. is transferred from Lygropia Lederer, 1863 based on characters in common with Conchylodes: 4.2.5. Udeini Mally, Hayden, Neinhuis, Jordal & upward-curled transtilla arms, white wings with spots Nuss trib.n. (orange in C. octonalis, black in congeners), corpus bur- sae with anterior appendix bursae, larvae feeding on Bor- Type genus: Udea Guenée, 1845 (in Duponchel) aginaceae (Powell & Opler 2009). Food plants. The food plant spectrum is broad in Udeini, Synapomorphies. 55:1, depth of gap/split of juxta being and several Udea species such as U. ferrugalis, U. lutea­ 10 – 60% of dorsoventral length of juxta; 99:1, strongly lis, U. olivalis, U. prunalis and U. rubigalis are pronounc- sclerotised colliculum anterior of the antrum and poste- edly polyphagous (Weigel et al. 1925; Lhomme 1935). rior of the attachment of the ductus seminalis present (not Mnesictena favidalis is recorded from Muehlenbeckia found with parsimony); 109:2, signum elongate rhombi- (Polygonaceae), M. notata from and Australina cal to ovate, longer than wide (found with parsimony). (Urticaceae) (Robinson et al. 2010). The larvae of Con­ Description. The uncus varies from unicapitate in the chylodes ovulalis (Guenée, 1854) are recorded to feed on Udea group (Deana Butler, 1879, Mnesictena Meyrick, Platanus (Platanaceae) (Solis 2008), other Conchylodes 1884, Tanaophysa Warren, 1892, Udea Guenée in Du- species feed on , Cordiaceae, Malvaceae, Bor- ponchel, 1845, Udeoides Maes, 2006) to conical (Con­ aginaceae and Annonaceae (Janzen & Hallwachs 2009). chylodes Guenée, 1854), reduced to triangle in Sisyrac­ Sisyracera and Cheverella are on Boraginaceae (Dyar era Möschler, 1890 and Ercta Walker, 1859, and reduced 1917; Wolcott 1950; Landry et al. 2011). to a transverse arching band in Cheverella Landry, 2011. Remarks. The genus Mnesictena was synonymised with The uncus dorsally has bifurcate chaetae in the Udea Udea by Munroe (1983), followed by Shaffer et al. group, but the chaetae are simple and located dorsally (1996). The type species of both genera were studied by and ventrally in Conchylodes, Sisyracera and Cheverel­ Mally & Nuss (2011) and found to be not congeneric, la, and lost in Ercta. The costa of the valva is slightly supporting Dugdale’s (1988) view of keeping them as concave; the ventral sacculus edge is parallel to the costa separate groups, but the authors did not reinstate Mne­

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Fig. 5. Udeini. A: adult female of Udea maderensis. B: adult male of . C: tympanal organs of male Udeoides mus­ cosalis. D: male genitalia of U. rhododendronalis, phallus omitted. E: male genitalia of C. zebra, phallus omitted. F: female genitalia of U. rhododendronalis. G: female genitalia of C. zebra. — Scale bars: A, B – 5 mm; C – G – 500 µm. sictena as bona genus. We leave this issue to a thorough the relationship of Cheverella, a Galápagos endemic, future study of the relationships within Udeini. prompted lengthy discussion in Landry et al. (2011), Sisyracera and Cheverella are problematic Neotropi- who decided that the Hydriris or Siga groups were the cal genera. Munroe (1995) left Sisyracera unplaced, and most likely places. Their relationship with Conchylodes

169 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

seems more plausible to us. The larvae of all three genera Niphopyralis (= Wurthia) (8 spp.) in Wurthiini. Further- feed on Boraginaceae. The moths are white with black more, based on morphological investigation, we place spotted lines (dense reticulate pattern in Sisyracera) and Diaphantania (3 spp.), Mimetebulea (1 sp.) and Pseude­ have ascending labial palpi. The valvae are attenuate, and bulea (4 spp.) in this tribe. This group can be considered the uncus is reduced or lost, bearing only fne chaetae. as an enlargement of Munroe’s (1995) Diaphantania ge- Sisyracera shares with Conchylodes an appendix bursae. nus group. The homoplasies to be accounted for are the change of Food plants. Food plants are not known for most Wurthi- the signum in Sisyracera and Cheverella (signum absent ini. The larvae of Apilocrocis glaucosia (Hampson, 1912) in the former, a small thorn in the latter) and loss of the feed on Celtis iguanea (Ulmaceae) (Janzen & Hallwachs valva fbula in Sisyracera. The robust valvae and infated 2009). Niphopyralis larvae live as brood parasites in nests sacculus of Cheverella can be explained as part of the of ants of the genera Oecophylla Smith, 1860 and Pol­ internally feeding larval syndrome, which is paralleled in yrhachis Smith, 1857 and feed on eggs, larvae, and pupae the Beebea group (Asciodini) and among the internally of their hosts (Roepke 1916; Kemner 1923). feeding Margaroniini that had been classifed in Mun- Remarks. The placement of Niphopyralis in Spilomeli- roe’s (1995) polyphyletic group. nae was a surprising discovery of Regier et al. (2012), Microphysetica Hampson, 1917 belongs to Hydririni but its particular association with Aristebulea and Api­ or Udeini. Females have a rhombiform signum and ap- locrocis in our analysis allows a radical but satisfying pendix bursae but not the tubular colliculum of Udeini. reinterpretation of the aberrant male genitalia (Fig. 6E). Males have sternite 8 like Udea and Choristostigma: The genitalia have been previously illustrated in Maes centrally membranous but without a distinctive anterior (1998a), who interpreted the gnathos as consisting of two scale feld, and the juxta is mesally weak but not split into separate, articulated arms. This condition occurs in other distinct arms. lepidopteran superfamilies (e.g. ), but it is not common in Pyraloidea (to our knowledge, occurring elsewhere only in Heliothelinae). In our interpretation, 4.2.6. Wurthiini + euspilomeline clades these two separate processes are the distal halves of the true valvae. They are small and displaced dorsad, but they Synapomorphies. 109:0, shape of anterior-most signum: have the same shape as the valvae in other wurthiines: circular, spinose, or invaginated as a spine, without obvi- distally attenuate with a triangular swelling at the base of ous axes. 103:1, indistinct division of the ductus bursae the costa (like that in Diaphantania impulsalis (Herrich- and corpus bursae (only found with parsimony). Schäffer, 1871) and Aristebulea principis). The valva of other Wurthiini genera is divided by a membranous cleft between the sacculus and distal half; this cleft reaches 4.2.7. Wurthiini Roepke, 1916 stat.rev. the outer margin in Apilocrocis and nearly so in the other genera. We interpret the sclerotised structures fanking Type genus: Wurthia Roepke, 1916 = Niphopyralis Hampson, 1893 the juxta in Niphopyalis as the valva sacculi, each with a median process similar to those in Aristebulea, Mimete­ Synapomorphies. 55:2, depth of gap/split of juxta rang- bulea, Pseudebulea, Diaphantania and Apilocrocis. It is ing from more than 60% of dorsoventral length of juxta not entirely clear whether the elongate processes in the to complete division into two juxta arms; 66:1, general ventral region of the genitalic arise from the sac- shape of fbula elongate, length at least twice the width, culi or from the juxta (as coded in the character list 55:2). apically rounded (may be curved). Slow optimization We further interpret the pair of weakly setose structures only: 24:0, venulae secundae absent. atop the tegumen to be the uncus in normal position, Description. The male genitalia have a broad triangu- even though the other members of the tribe have a single- lar, ventrally directed fbula; the mesal sides of the sac- headed uncus with bifd chaetae. The gnathos is absent. culi are produced as two strongly sclerotized arms which dorsally end in a broad, spinulose tip or a slim, needle- shaped projection (not split in Mimetebulea Munroe & 4.2.8. Euspilomeline clades Mutuura, 1968); the mediodorsal sacculus has a medially directed process (absent in Apilocrocis Amsel, 1956 and Synapomorphies. 86:1, reduction of phallus apodeme to Diaphantania Möschler, 1890). In the female genitalia, a ventral, longitudinally sclerotized strip along the man- the lamella antevaginalis forms a strongly sclerotised an- ica (the rest of the apodeme being more or less membra- trum frame; the signum is rounded, small (Aristebulea nous). In addition, the parsimony trees add many more Munroe & Mutuura, 1968, Pseudebulea Butler, 1881) to synapomorphies: 32:1, male tergite 8 with anterior edge relatively large (Apilocrocis, Diaphantania), and absent emarginate; 60:1, partly sclerotized hair pencils present in Mimetebulea and Niphopyralis. Under fast optimiza- on anterior edge of vinculum-tegumen connection; 78:2, tion, the loss of venulae secundae (24:0) is shared with valva with convex costa; 84:0, phallus without coecum; various Hydririni, Lineodini and Udeini. 95:1, antrum with longitudinal membranous strip; and Systematics. Based on our phylogenetic results, we 99:0, a strongly sclerotised colliculum between antrum place Apilocrocis (11 spp.), Aristebulea (2 spp.) and and ductus seminalis absent.

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Fig. 6. Wurthiini. A: head of female Niphopyralis sp., ventral view. B: head of female Niphopyralis sp., dorsal view. C: tympanal organs of male Niphopyralis sp. D: adult female of Niphopyralis sp., posterior abdomen removed. E: male genitalia of Niphopyralis sp. F: male genitalia of Apilocrocis novateutonialis, phallus omitted. G: posterior abdomen of male N. chionesis. H: female genitalia of Diaphantania impulsalis. I: female genitalia of N. chionesis. — Scale bars: D – 5 mm; F – I – 500 µm.

171 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

In this monophylum, the postmedial (PM) lines of The placement of Pycnarmon among the euspilomeline both wings are usually more jagged than in Pyraustinae clades is still uncertain. Based on morphological charac- and among the early-diverging spilomeline clades. Es- teristics, we further place the following genera in Agro- pecially the hindwing PM line is usually boldly marked terini: Munroe, 1974a (2 spp.), Bocchoropsis Am- and projects distad on the M and CuA veins, whereas in sel, 1956 (2 spp.), Chalcidoptera Butler, 1887 (15 spp.), Pyraustinae, the hindwing PM line tends to be faint and Chilochromopsis Munroe, 1964 (1 sp.), Coenostolopsis a smooth arc. In Pyraustinae, only very rarely does the Munroe, 1960 (3 spp.), Hübner, 1818 (12 spp.), forewing PM line jut basad on the anal fold or is drawn Framinghamia (2 spp.), Glaucobotys Maes, 2008 (1 sp.), costad toward the discal spot (but see Pseudopyrausta). Goliathodes Munroe, 1974a (1 sp.), Gypodes Munroe, In general, the hindwing PM line is similar to the fore- 1976 (1 sp.), Lygropia Lederer, 1863 (68 spp.), Lypoti­ wing PM line, so the combination of boldness and jag- gris Hübner, 1825 (1 sp.), Micromartinia Amsel, 1957 (1 gedness distinguishes it. Although the jagged shape is sp.), Microthyris Lederer, 1863 (7 spp.), Nagiella Mun- common in the euspilomeline clade, it is also present in roe, 1976 (4 spp.), Lederer, 1863 (26 spp.), Aristebulea and Pseudebulea (Wurthiini). Meyrick, 1884 (18 spp.), Leder- er, 1863 (9 spp.), Phaedropsis Warren, 1890 (24 spp.), Guenée, 1854 (26 spp.), Tetracona Mey- 4.2.9. Agroterini Acloque, 1897 stat.rev. rick, 1884 (2 spp.) and Ulopeza Zeller, 1852 (16 spp.). Nagiella has been considered either a valid genus Type genus: Agrotera Schrank, 1802 (Munroe 1976b; Kirti & Sodhi 2001; Rose 2001; Ul- lah et al. 2017) or a synonym of Pleuroptya (= Patania) Synapomorphies. 47:0, structure of uncus head chaetae (Inoue 1982; Leraut 1997). We concur with Munroe’s simple, not split; 58:1, ratio between saccus length and (1976b) separation of Nagiella from Patania. Ullah et sacculus breadth > 1 (elongate saccus, often with bul- al. (2017) describe a fourth species in this genus. For the bous apex). The parsimony trees add 8:0, sclerotization generic diagnosis see Munroe (1976b). on vesica absent; and 44:0, conical (non-capitate) uncus. This diverse, globally distributed tribe generally cor- Slow optimization only: 8:0, 3rd labial palpomere direct- responds to Munroe’s (1995) Syllepte group. We conjec- ed dorsally. ture that he placed the Phaedropsis and Syllepte groups Description. The labial palps are upturned. The uncus frst in his checklist because some have a gnathos in the has a broad base, usually truncate to variously reduced, traditional sense, i.e. with a well-developed medial pro- at the extreme being a squat, transversely rectangular cess, which would seem to be the primitive state. In our square. The gnathos has a well-developed medial process analysis, this process is secondarily derived, since none in some genera (see e.g. Leraut 2005b: fgs. 14 – 17). of the non-euspilomeline clades have it (the gnathos be- The valvae are more or less rectangular, with costal and ing a simple, transverse band). This process is also pre- ventral margins parallel, or slightly ovate; the saccus sent in two genera of uncertain placement: Mimudea is notably elongate, often distally bulbous. The female Warren, 1892, and Deuterophysa Warren, 1889. genitalia have round and granular signa, single or double, Species of Phaedropsis are hardly separable from the rarely extended as horns (Framinghamia Strand, 1920, type species of Lygropia, Asopia unicoloralis Guenée, Phostria oajacalis (Walker, 1866)). 1854. Lygropia and Phostria are major dustbin genera The upturned 3rd labial palpomere is a synapomor- of this tribe, holding many explictly misplaced species phy for this tribe in the DELTRAN analysis, but it is (Munroe 1995). shared with various other tribes, such as Asciodini, the In many genera, especially in the Old World, the fore- Siga group of Spilomelini, Spoladea, and some Nomo- wing costa bears a light-colored triangular spot. This is philini and Steniini. The presence of simple, unsplit the “Nosophora-Chalcidoptera” group referred to by uncus chaetae is paralleled in some Pyraustinae, Marga- Munroe (1974a). In some taxa, the spot is so strongly roniini and Udeini, and in Nomophila Hübner, 1825. In developed that it extends to the tornus and flls most of many genera, the tegumen mesally extends anteriad, like the forewing (e.g. some misplaced ‘Leucinodes’ species, an extended roof. This unique structure may characterize ‘Syllepte’ dottoalis Schaus, 1927). a clade in Agroterini. In some genera, the papillae anales Food plants. Larvae are generally leaf-tiers. Larvae of face ventrad at a right angle to the axis of the ovipositor; Patania silicalis and P. sabinusalis have been reared this state is paralleled in some Margaroniini. on Urticaceae (Kimball 1965; Miller et al. 2007; So- Systematics. According to our phylogenetic results, lis 2008), P. silicalis furthermore on Polygonum (Po- Agroterini comprises Aetholix Lederer, 1863 (4 spp.), lygonaceae), Ipomoea and Merremia (Convolvulaceae), Agrotera (27 spp.), Haritalodes Warren, 1890 (11 spp.), Rivina (Petiveriaceae) and Bougainvillea (Nyctaginace- Neoanalthes Yamanaka & Kirpichnikova, 1993 (8 spp.), ae) (Heppner & Habeck 1976; Bendicho-Lopez 1998); Patania Moore, 1888 (= Pleuroptya Meyrick, 1890) (41 P. ruralis feeds on Urtica (Urticaceae), Humulus (Can- spp.), Phostria Hübner, 1819 (87 spp.) and ‘Pycnarmon’ nabaceae), Chenopodium, Atriplex (), pantherata Butler, 1878 which is not congeneric with Filipendula (Rosaceae) and Ribes (Grossulariaceae) P. jaguaralis (Guenée, 1854), the type species of the (Lhomme 1935); Central American Patania species (as polyphyletic genus Pycnarmon Lederer, 1863 (59 spp.). Pleuroptya) are recorded from Acanthaceae,

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Fig. 7. Agroterini. A: adult male of Notarcha cf. quaternalis. B: adult male of Pycnarmon pantherata, abdomen removed. C: male genita- lia of P. pantherata, phallus omitted. D: male genitalia of Agrotera nemoralis. E: female genitalia of A. nemoralis. F: phallus of Patania ruralis. G: tympanal organs of male Phostria temira. — Scale bars: A, B – 5 mm; C–G – 500 µm.

and Urticaceae (Janzen & Hallwachs 2009). Known Signifcant host associations are with Malvaceae food plants for Phostria larvae are mainly Convolvu- s.l. (Pantographa, Haritalodes, Phaedropsis), Convol- laceae, Malvaceae and Rubiaceae (Janzen & Hallwachs vulaceae (Phostria tedea-group, Lygropia tripunctata- 2009). Agrotera nemoralis feeds on Carpinus, Betula, group, Microthyris incl. Cyclocena; see Hayden & Dickel Corylus (), Castanea and Quercus () 2014) and Rubiaceae (‘’ xanthozonalis-group). (Melzer & Nuss 2009), while two Australasian Agrotera Remarks. The mimetic ‘Pilocrocis’ xanthozonalis-group species are reported to feed on Syzygium spp. (Myrta- belongs here, and its species are misplaced in the Herpe­ ­ ceae) (Miller et al. 2007). Haritalodes is recorded from togrammatini genus Pilocrocis. Malvaceae, Amaranthaceae and Moraceae (Ghesquiére The Australian species of Agrotera have recently been 1942; Miller et al. 2007); Diastictis on Asteraceae (Pow- revised by Chen et al. (2017), who removed Leucinodella ell & Opler 2009); Framinghamia on Salix (Salicaceae); Strand, 1918, Nistra Walker, 1859, Sagariphora Mey- Phaedropsis on Polygonaceae and Malvaceae (Janzen & rick, 1894 and Tetracona from synonymy with Agrotera; Hallwachs 2009). the former three genera can currently not be placed in any

173 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

of the proposed tribes, but Tetracona is placed in Agro- 1859), Samea ecclesialis Guenée, 1854 and Prophantis terini, close to Aetholix (Chen et al. 2017). xanthomeralis Hampson, 1918 comb.n.). Systematics. Based on our phylogenetic fndings we as- sociate these taxa with Margaroniini: Agathodes Guenée, 4.2.10. Margaroniini Swinhoe & Cotes, 1889 1854 (16 spp.), Agrioglypta Meyrick, 1932 (11 spp.), stat.rev. Antigastra Lederer, 1863 (2 spp.), Asturodes Amsel, 1956 (1 sp.), Azochis Walker, 1859 (16 spp.), Botyodes Type genus: Margaronia Hübner, 1825 = Dichocrociinae Swinhoe, 1900: 478 Guenée, 1854 (10 spp.), Cadarena Moore, 1886 (1 sp.), = Hapaliadae Swinhoe, 1890: 268 Conogethes pandamalis (Walker, 1859) comb.n., Cyda­ = Margarodidae Guenée, 1854: 286 lima Lederer, 1863 (9 spp.), Diaphania Hübner, 1818 (95 spp.), Dichocrocis cf. zebralis (Moore, 1867), Filodes Synapomorphies. 33:1, sclerotization of male tergite Guenée, 1854 (16 spp.), Ghesquierellana Berger, 1955 8: heterogenous, i.e. with distinct sclerotization pattern; (5 spp.), Glyphodes Guenée, 1854 (156 spp.), Hodeber­ 72:1, extension (process in some cases) at dorsodistal tia Leraut, 2003 (1 sp.), Liopasia (15 spp.), Leucochroma sacculus present (not found with parsimony); 75:1, joint Guenée, 1854 (6 spp.), Maruca Walker, 1859 (4 spp.), of basal valva costa (with vinculum) extended into an Megastes (16 spp.), ‘Nacoleia’ insolitalis (Walker, 1862), elongate, rod-shaped process present; 103:0, demarca- Obtusipalpis (6 spp.), Omiodes Guenée, 1854 (98 spp.), tion between corpus bursae and ductus bursae distinct, Palpita Hübner, 1808 (162 spp.), Prenesta Snellen, 1875 with a narrow anterior ductus transforming into a wide (18 spp.), Pygospila Guenée, 1854 (10 spp.), Rhimphalea corpus bursae (only found with parsimony); 108:1, two Lederer, 1863 (12 spp.), Terastia Guenée, 1854 (7 spp.), signa. Slow optimization only: 40:1, sclerite on each Zebronia Hübner, 1821 (6 spp.). ‘Nacoleia’ insolitalis is pleural membrane of male segment 8 present (not found misplaced in Nacoleia Walker, 1859, and its correct ge- with parsimony); 95:1, longitudinal membranous strip in neric affliation remains uncertain. ‘Dichocrocis’ panda­ the antrum sclerotisation present (not found with parsi- is misplaced in Dichocrocis; its correct placement mony). is in Conogethes Meyrick, 1884, where it is transferred Description. The adult moths are mostly medium-sized here (see above). Dichocrocis Lederer, 1863 (53 spp.) is to large. Many genera have the scape and pedicel of the considered polyphyletic and needs revision; maculation male antenna modifed. Males of many genera have a and male genitalia of the type species D. frenatalis Leder- large, black tuft of fne, hairlike sex-scales on A8. The er, 1863 indicate a placement among the euspilomeline tegumen is often spacious in sagittal dimension (hamper- groups, probably near or in Steniini, but this needs further ing the planar mounting of the genitalia on a glass slide). investigation. The uncus is conical or unicapitate and long-necked (bi- ‘Glyphodes’ rubrocinctalis is misplaced in Glyphodes; furcate in Cydalima perspectalis), with simple or bifur- in our phylogenetic analysis (Fig. 1) it is subordinate in cate chaetae, often with both, or without chaetae. The Prenesta. The male genitalia are smaller than those of anterior tegumen-vinculum connection usually has an at- the type species of Prenesta, P. scyllalis, but they are tached pad of hairpencils, the pad structure ranging from similar in structure, and the moths share distinctive red a simple sclerotized base with one kind of simple hairs to and yellow maculation. We therefore transfer Prenesta several membrane-connected sclerites with several dif- rubrocinctalis (Guenée, 1854) comb.n. from Glyphodes. ferently structured hairs. The valva is commonly broad Furthermore, we assign the following taxa based and oval, with one fbula about halfway to the apex. The on morphological investigation: Alytana J.C. Shaffer sacculus is comma-shaped, broadest at the anteroventral & Munroe, 2007 (2 sp.), Anarmodia Lederer, 1863 (24 valva base, arching and tapering distad (broadening in spp.), Aphytoceros Meyrick, 1884 (3 spp.), Arthroschis­ Liopasia and Obtusipalpis), its distal apex usually ending ta Hampson, 1893 (2 spp.), Caprinia Walker, 1859 (11 in a ridge or sclerotized process in close spatial associa- spp.), Chabulina J.C. Shaffer, & Munroe, 2007 (2 spp.), tion with the distal fbula. The vesica of the phallus has Charitoprepes Warren, 1896 (2 sp.), Chrysophyllis Mey- a granulated area and/or one to several bodkin-shaped rick, 1934 (1 sp.), Chrysothyridia Munroe, 1967 (2 spp.), cornuti. The corpus bursae often has a pair of circular Lederer, 1863 (38 spp.), Colomychus signa, which can be fat or invaginated to form spikes, Munroe, 1956 (2 spp.), Amsel, 1956 (4 spp.), or signum absent. The ductus bursae is granular in many Lederer, 1863 (4 spp.), Conogethes (16 genera (e.g. Cydalima). spp.), Didymostoma Warren, 1892 (2 spp.), Dysallacta The heterogenous sclerotization of male tergite 8 Lederer, 1863 (3 spp.), Endocrossis Meyrick, 1889 (4 (33:0) may be shared also with Asciodini, Spoladea, spp.), Eusabena Snellen, 1901 (4 spp.), Glyphodella J.C. Trichaeini, and some Steniini. Shaffer, & Munroe, 2007 (3 spp.), Hedyleptopsis Mun- The base of the valva costa simple, not rod-shaped, in roe, 1960 (1 sp.), Heterocnephes Lederer, 1863 (4 spp.), several genera: the stout-bodied Liopasia Möschler, 1882, Hoterodes Guenée, 1854 (5 spp.), Loxmaionia Schaus, Megastes Guenée, 1854 and Obtusipalpis Hampson, 1913 (1 sp.), Marwitzia Gaede, 1917 (3 spp.), Mega­ 1896, and in ‘Glyphodes’ rubrocinctalis (Guenée, 1854) physa Guenée, 1854 (1 sp.), Meroctena Lederer, 1863 (4 and Zebronia phenice (Stoll, 1782). The rod-shaped state spp.), Nolckenia Snellen, 1875 (1 sp.), Moore, is paralleled in Hymeniini, Arthromastix lauralis (Walker, 1886 (10 spp.), Pachynoa Lederer, 1863 (12 spp.), Paro­

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Fig. 8. Margaroniini. A: tympanal organs of female Cydalima perspectalis. B: left valva of male genitalia of Conogethes pandamalis. C: phallus of male C. pandamalis. D: male adult of Glyphodes prothymalis. E: 7th sternite of male Azochis cf. rufdiscalis. F: head of male Azochis sp. G: 7th abdominal sternite of male Rhimphalea cf. astrigalis. H: dissected abdomen of male Terastia meticulosalis. I: ventral wing side of male G. prothymalis with frenulum bristle. J: female genitalia of C. perspectalis. K: female genitalia of Omiodes continuata­ lis. — Scale bars: A – C, E, G, H, J, K – 500 µm; D – 5 mm.

175 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

tis Hübner, 1831 (37 spp.), Poliobotys J.C. Shaffer & 1782) are leaf-tiers on Lamiales: The former, best known Munroe, 2007 (1 sp.), Polygrammodes Guenée, 1854 (78 as a pest of (), also feeds on Bigno- spp.), Polygrammopsis Munroe, 1960 (1 sp.), Radessa niaceae (Tecoma stans) and Plantaginaceae (Powell & Munroe, 1977 (2 spp.), Rhagoba Moore, 1888 (2 spp.), Opler 2009). The latter feeds on Bignoniaceae but was Sinomphisa Munroe, 1958 (3 spp.), Sparagmia Guenée, also recorded on (Malvaceae) and Ricinus 1854 (1 sp.), Lederer, 1863 (8 spp.), Syn­ (Euphorbiaceae) (Robinson et al. 2010). Hodebertia clera Lederer, 1863 (13 spp.), Syngamilyta Strand, 1920 testalis (Fabricius, 1794) larvae predominantly feed on (5 spp.), Moore, 1885 (9 spp.), Tessema J.F.G. Asclepiadaceae (Robinson et al. 2010). Botyodes feeds Clarke, 1986 (1 sp.), Tyspanodes Warren, 1891 (20 spp.), on Flacourtiaceae, Moraceae, Salix (Salicaceae) and sev- Uncobotyodes Kirti & Rose, 1990 (1 sp.). eral other hosts (Nakamura & Ohgushi 2004; Robinson Tyspanodes is not a natural group, and at least T. ex­ et al. 2010). Cadarena pudoraria (Hübner, 1825) and althealis (Walker, 1859) is misplaced here; we did not in- the closely related ‘Glyphodes’ (or ‘Pyrausta’) perel­ vestigate the type species, T. nigrolinealis (Moore, 1867), egans (Hampson, 1898) group are recorded from Passi- but we can confdently place T. hillalis Schaus, 1927, T. foraceae, C. pudoraria also from Gossypium and hypsalis Warren, 1891 and T. celebensis Munroe, 1960 in (Malvaceae) (Janzen & Hallwachs 2009; Robinson et al. Margaroniini. 2010; de Prins & Mazzei 2016). Ghesquierellana hirtu­ Heterocnephes apicipicta Inoue, 1963 is misplaced salis (Walker, 1859) larvae feed on Ficus (Moraceae) and in Heterocnephes and transferred to the monotypic Gossypium (Malvaceae); Megastes on Ipomoea (Convol- Charitoprepes as Charitoprepes apicipicta (Inoue, vulaceae); Azochis on Ficus (Moraceae); Conogethes lar- 1963) comb.n. The type species C. lubricosa Warren, vae are recorded from a wide range of plants, e.g. Pinace- 1896 shares with C. apicipicta the wing pattern and the ae, Gnetaceae, Malvaceae, Sapindaceae, Euphorbiaceae structure of the male genitalia (cf. Inoue 1963; Kim et al. and Zingiberaceae (Robinson et al. 2010; Shashank et al. 2014). Alytana calligrammalis (Mabille, 1879) comb.n. 2018). ‘Nacoleia’ insolitalis from Sandoricum (Meliace- is transferred from Analyta, a transfer that had not been ae) (Robinson et al. 2010). The known larval host plants formerly proposed by Shaffer & Munroe (2007) in their of the Glyphodes genus group sensu Sutrisno (2002b) description of Alytana. (Glyphodes, Dysallacta, Talanga, Agrioglypta) are pri- Food plants. We arrange food plant records, where avail- marily the latex-containing Moraceae and Apocynaceae able, according to the clades found within Margaroniini (Kimball 1965; Common 1990; Robinson et al. 2010). as shown in Fig. 1: Astrodes fmbriauralis is recorded Obtusipalpis is recorded from Rubiaceae, Moraceae and from Colubrina (); Maruca vitrata is a pest Rutaceae (Robinson et al. 2010). species on various Fabaceae such as Lablab, Phaseolus, Host plants for other genera placed in Margaroniini Pisum, Psophocarpus, Sesbania and Vigna, but has also are: and on Rubiaceae, the latter been recorded from Rubiaceae, Solanaceae, and also on Apocynaceae, on which Pygospila and Stemor­ Euphorbiaceae (Robinson et al. 2010). Cydalima mainly rhages mainly feed; Cirrhochrista on Moraceae; Con­ feeds on Apocynaceae (C. laticostalis (Guenée, 1854)), dylorrhiza on Salicaceae; Eusabena on Hoya (Asclepia- Buxaceae (C. perspectalis (Walker, 1859)) or Rhamnace- daceae); Anarmodia and Sparagmia on Araliaceae; Syn­ ae (C. mysteris Meyrick, 1886) (Robinson et al. 2010). clera on Gouania and Zizyphus (Rhamnaceae) (Mann & Filodes feeds on Thunbergia (Acanthaceae) (Robinson Brar 1980; Janzen & Hallwachs 2009; Robinson et al. et al. 2010). Diaphania species mostly feed on Cucur- 2010; Hayden et al. 2017). bitaceae; Palpita is primarily on Oleaceae, but P. fegia, The most general trend in this group is feeding on la- probably the frst-diverging member with a plesiomor- tex-bearing plants, especially Apocynaceae and Morace- phic male antennal scape, feeds on Apocynaceae (Moore ae. The habit of boring in tubers of Ipomoea (Convol- 1884 – 1887; Hinckley 1964; Kimball 1965; Clavijo Al- vulaceae) by Megastes, Polygrammodes eleuata, and bertos 1990; Solis 2006, 2008; Robinson et al. 2010). (Guenée, 1854) is explained by Omiodes species feed on a variety of host plants, and two the presence of latex in these roots. Nevertheless, many species (O. diemenalis (Guenée, 1854), O. indicata (Fab- genera and genus groups diverge from the pattern and ricius, 1775)) are widespread pests on Fabaceae; the lar- radiate on non-latex-bearing plants, e.g. some Polygram­ vae of the Hawaiian Omiodes clade feed on monocotyle- modes in roots of Vernonia, Sinomphisa in Bignoniaceae, donous plants, except O. monogona Moore, 1888, which and (Hampson, 1896) in bamboo. feeds on Fabaceae (Robinson et al. 2010; Haines & Ru- Remarks. Margaroniini roughly refects a combination binoff 2012). Omiodes stigmosalis Warren, 1892, a borer of Munroe’s (1995) Diaphania and Polygrammodes in fg fruits (Janzen & Hallwachs 2009), is misplaced in groups. The assumed close relationship for Agathodes, Omiodes but has the characters of Margaroniini. Pren­ Terastia and Liopasia (Munroe 1960; Sourakov et al. esta is recorded mainly from Apocynaceae and Moraceae 2015) has been confrmed by our results. Females of the (Janzen & Hallwachs 2009). Larvae of Liopasia, Ag­ Agathodes genus group exhibit an appendix bursae (ab- athodes and Terastia commonly feed on Erythrina (Fa- sent in some species); the larvae are feeding on Erythrina baceae) (Hinckley 1864; Kimball 1965; Sourakov 2012; (Fabaceae) (Pereira et al. 2014). Pereira et al. 2014). Antigastra catalaunalis (Duponche- The possible sister group relationship between Omi­ lia, 1833) and Zebronia phenice (Stoll in Cramer & Stoll, odes and Cnaphalocrocis Lederer, 1863, as suggested by

176 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Haines & Rubinoff (2012), could not be confrmed. We the uncus entirely split into two separate unci (unsplit in treat Cnaphalocrocis as a member of Spilomelini (see Zeuzerobotys), bearing bifd chaetae; the costal margin of below), whereas Omiodes belongs to Margaroniini. The the valva is approximately straight or only slightly con- sister group of Omiodes remains to be discovered. vex near the base, never strongly convex, distally straight clavijo Albertos (1990) observed and described the or slightly concave; the apical half of the valva is bluntly “anepisternal scale organ” in males of many genera of attenuate, slightly to markedly narrower than basal half Spilomelinae. Apparently, this is a tymbal organ (Naka- of valva with its infated sacculus; there is one ventrally no et al. 2012b). Among Spilomelinae, ultrasound pro- directed, hook-shaped to spatulate fbula emerging from duction is reported from the Margaroniini Conogethes centre of valva. The combination of the bifd uncus and punctiferalis (Guenée, 1854), Walk- the shape of the valva distinguish members of the Siga er, 1859 and (Bremer, 1864) as group from robust-bodied Margaroniini. The ductus bur- well as in Spoladea recurvalis (Fabricius, 1775) (Hyme- sae is as long as or shorter than corpus bursae, and the niini) (Nakano et al. 2009, 2012a). corpus bursae is spherical, rarely ovate, without signa. Systematics. Based on our phylogenetic analyses, we place the following taxa in Spilomelini: Cnaphalocrocis 4.2.11. Spilomelini Guenée, 1854 stat.rev. group with Cnaphalocrocis (27 spp.), Marasmia Lederer, Type genus: Spilomela Guenée, 1854 1863 (9 spp.), Salbia Guenée, 1854 (35 spp.) and Spilo­ = Siginae Hampson, 1918 mela Guenée, 1854 (8 spp.); Siga group with Eporidia Walker, 1859 (1 sp.) and Siga Hübner, 1820 (2 spp.), as Synapomorphies. 45:2, apical uncus bifurcate. The un- presumed by Munroe (1958). cus is bicapitate with bulbous heads. Based on morphological investigation, we further as- Description. Small (15 mm wingspan) to large (90 mm sign Aethaloessa Lederer, 1863 (3 spp.), Geshna Dyar, wingspan) moths. Spilomelini shares the bicapitate uncus 1906 (1 sp.), Marasmianympha Munroe, 1991 (1 sp.), with Asciodini and some Steniini such as Metasia and Orphanostigma Warren, 1890 (6 spp.), Palpusia Am- Loxostegopsis. The costa of the valva is straight to slight- sel, 1956 (10 spp.) and Rhectocraspeda Warren, 1892 (2 ly concave or convex. This tribe consists of two distinct spp.) to the Cnaphalocrocis group, and Cirrhocephalina clades: the Cnaphalocrocis group and the Siga group. Munroe, 1995 (5 spp.), Scaptesylodes Munroe, 1976 (2 In the Cnaphalocrocis group, adults are small to spp.) and Zeuzerobotys Munroe, 1963 (1 sp.) to the Siga medium-sized. Most genera are brown and drab in col- group, following Munroe (1963; 1976b; 1995) and Lan- our, whereas Spilomela has contrasting maculation; dry et al. (2011) (but see Remarks). the wings’ transverse lines consist of straight segments Food plants. The hosts of the Cnaphalocrocis group are and angulate junctions. Males have a feld of enlarged, heterogeneous, but the group includes a major radiation raised scales on the centre of the forewing costa (absent on monocots, especially on Gramineae. The larvae of in several taxa). Sacci tympani are small and closely set. Aethaloessa, Cnaphalocrocis, Marasmia and Salbia are In the male genitalia, the fattened uncus is weakly bifd, mainly leaf-rollers on Poaceae, Salbia larvae are also re- the head consisting of two connected, fat pads or felds corded from Verbenaceae and to a lesser amount from of chaetae rather than clearly separate parts; the uncus is Gesneriaceae and Fabaceae, and Aethaloessa foridalis lost in Geshna. The sacculus ends distally with a small (Zeller, 1852) from Urticaceae (Janzen & Hallwachs fbula-like process pointing inward toward the center of 2009; Robinson et al. 2010). Rhectocraspeda is found the valva. Some taxa have a fbula in the center of the on Piper (Piperaceae), Columnea (Gesneriaceae) and valva. In Spilomela perspicata, the details of the male Solanaceae, Geshna on (Cannaceae), Lilium (Lil- genitalia are distorted by elongation, but the uncus is iaceae), Thalia geniculata (Marantaceae) and Zantede­ apically bifd on close inspection. In females, the col- schia (Araceae), Palpusia on Convolvulaceae and Ru- liculum is cylindrical and open dorsally or entire. The biaceae, and Orphanostigma on Lamiaceae, Asteraceae colliculum is often extended as extra sclerotization on and Malvaceae (Kimball 1965; Heppner 2003; Janzen & the adjacent ductus bursae. The ductus bursae is usually Hallwachs 2009; Robinson et al. 2010). Spilomela larvae very short and has fne spinules or striations next to the are reported from Dilleniaceae, Ulmaceae and Rubiaceae colliculum. The signum typically is a granulose circle (Janzen & Hallwachs 2009). or a small thorn, but Palpusia species have two long, In the Siga group, the hosts are unknown for Siga and sickle-shaped signa, and Spilomela receptalis (Walker, Eporidia, the two genera included in our phylogenetic 1859) has two shorter sickles; signa are absent in Rhec­ analysis. tocraspeda and Spilomela perspicata (Fabricius, 1787) Remarks. The genus Spilomela is polyphyletic and itself, which in addition has a very long, unsclerotized needs revision. We base our conclusions on the type spe- ductus bursae. cies, S. perspicata. The Siga group includes medium-sized to large and The Siga group shares with Asciodini a similar mor- thick-bodied moths. The proboscis is lost in Siga, other- phology of the male genitalia, especially the overall ro- wise normally developed. The sacci tympani are exposed bust form and bifd uncus. We transfer several genera to as a shallow zona glabra, and the fornix tympani is circu- Asciodini (below). Together with Siga and Eporidia, the larly rounded without an angle. The male genitalia have genera that we retain in the Siga group on morphologi-

177 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 9. Spilomelini. A: adult male of Spilomela perspicata. B: adult female of Marasmia poeyalis. C: adult female of Eporidia dariusalis, abdomen removed. D: 8th abdominal segment of male S. perspicata. E: male genitalia of S. perspicata, uncus detached from tegumen, phallus omitted. F: male genitalia of M. poeyalis, valvae embedded inverted in the preparation with the costa facing outward, phallus omit- ted. G: male genitalia of E. dariusalis, phallus omitted. H: male genitalia of Salbia cf. haemorrhoidalis, one valva and phallus omitted. I: uncus heads of the male genitalia of M. poeyalis. J: female genitalia of E. dariusalis. — Scale bars: A – C (same scale) – 5 mm; D – H, J – 500 µm; I – 100 µm.

178 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

cal grounds (Cirrhocephalina, Scaptesylodes, and Zeu­ elongate signum (Hileithia spp.); the signum is absent or zerobotys) share the same shape of fbula and valva, and rudimentary in Eurrhyparodes, but with posterior wall of non-infated transtilla. The colliculum of Siga, Eporidia, corpus bursae sclerotized; the corpus bursae is well dis- and Scaptesylodes is entire and bulges ventrad rather like tinguished from the long, slender, membranous or partly a pot-belly or a pitcher plant (Nepenthes) (female genita- sclerotized ductus bursae; the colliculum is membranous lia not observed for Cirrhocephalina and Zeuzerobotys). or with a sclerotisation partially encompassing the duc- The loss of the proboscis in Siga is paralleled in Wurthi- tus; the antrum is weakly to strongly sclerotized, simple. ini: Niphopyralis. Systematics. Based on our phylogenetic results, we place The monotypic Gesha is included because the larvae Eurrhyparodes Snellen, 1880 (12 spp.), Herpetogramma feed on Canna, the wing pattern is typical of the Cnaph­ Lederer, 1863 (100 spp.) and Hileithia Snellen, 1875 (19 alocrocis group, and the genitalia share characters with spp.) here. Furthermore, Lederer, 1863 some Salbia species, namely swellings on the base of the (85 spp.), Cryptobotys Munroe, 1956 (2 spp.) and Pi­ valva costa (also in S. mizaralis (Druce, 1899)) and the locrocis Lederer, 1863 (65 spp.) are assigned to Herpe- broad, triangular uncus. The loss of bifd uncus chaetae togrammatini based on morphological characters. and the movement of the fbula to a central position on Munroe (1995) further places the monotypic Pelinop­ the valva are homoplasies. sis Dognin, 1905 in his Herpetogramma group. As we did not study this taxon, we keep it unplaced. Food plants. The known food spectrum of the larvae 4.2.12. Herpetogrammatini Mally, Hayden, comprises Acanthaceae (Hileithia, Pilocrocis, Eurrhyp­ Neinhuis, Jordal & Nuss trib.n. arodes splendens Druce, 1895), Actinidiaceae (Pilocro­ cis), Malvaceae (Hileithia) and Urticaceae (Pilocrocis) Type genus: Herpetogramma Lederer, 1863 (Heppner 2003; Solis 2008; Janzen & Hallwachs 2009). ‘Pilocrocis’ milvinalis (Swinhoe, 1886) is reported from Synapomorphies. 5:1, length of sensillar setae at basal Apocynaceae, Fabaceae and Rubiaceae, P. pterygo­ antennomeres relative to diameter of basal antennomeres dia Hampson, 1912 from Lamiaceae (Robinson et al. in male > 50%; 107:0, corpus bursae sclerotisation con- 2010). Eurrhyparodes bracteolalis (Zeller, 1852) is re- sisting of a granulose area. Slow optimization only: 8:1, corded from Solanum (Solanaceae) and Oryza (Poaceae) 3rd labial palpomere porrect. The parsimony trees im- (Robinson et al. 2010), from ply one apomorphy: 44:0, a conical, non-captitate uncus; Chenopodium (Amaranthaceae) (Solis 2008). The spe- characters 5:1 and 107:0 are synapomorphies with As- cies-rich genus Herpetogramma (100 spp.; Nuss et al. ciodini. 2003 – 2019) contains species with a variety of food Description. The uncus is conical, non-capitate, slen- plants ranging from ferns to angiosperms (Solis 2008; der to broad, with dorsally attached chaetae that are Janzen & Hallwachs 2009). bifd in Eurrhyparodes and Hileithia and hairlike in all Remarks. The mimetic ‘Pilocrocis’ xanthozonalis other genera examined; the uncus is broadly attached Hampson, 1912 group (including P. cyrisalis (Druce, to the tegumen, that is, the tegumen grades evenly into 1895)) feeds on Rubiaceae; this group is misplaced in Pi­ the uncus without “shoulders”. Character 44:0, shape of locrocis and belongs to Agroterini, based on the extended uncus conical, non-capitate, is absent in several species tegumen, naked uncus, and twin tack-shaped signa. of Blepharomastix such as B. ranalis (Guenée, 1854), an apparent reversal of the tribe’s synapomorphy. The val- vae are ovate with a convex to straight costa and rounded 4.2.13. Hymeniini + Asciodini to acute apex; the sacculus is weakly developed or ab- sent; the fbula is long and emerges from near costa base, Synapomorphies. 61:1, two or more hairpencil sclerites pointing towards the centre of the ventral valva edge, or on each side of the genitalia (articulated with each other the fbula is reduced to a fold or lost entirely (Crypto­ via membranes); 62:1, more than one kind of hairpencil botys, Pilocrocis, some Herpetogramma spp.); the juxta chaetae present. is compact, dorsally split; the saccus is V-shaped, its tip often somewhat offset; the hairpencils are simple (one sclerite bearing one kind of simple chaetae) or absent. 4.2.14. Hymeniini Swinhoe, 1900 stat.rev. The phallus coecum is short or absent; the phallus ap- odeme is membranous apart from a ventral longitudinal Type genus: Hymenia Hübner, 1825 sclerotized strip; the vesica is granulose, often with a dense patch of small cornuti. The corpus bursae is mem- Synapomorphies. 3:1, transverse rim on anterior or me- branous or posteriorly with a granulose area, and the sal face of pedicellus in male present; 9:0, size of 3rd signum is single or absent: when most developed, it is labial palpomere well developed in both sexes; 75:1, a round, granulose rhomboid with transverse axis domi- joint of basal valva costa (with vinculum) extended into nant (Cryptobotys, Herpetogramma spp.), or reduced to an elongate, rod-shaped process. Slow optimization only: a transverse line (Pilocrocis ramentalis Lederer, 1863, 40:1, sclerite present on each pleural membrane of male Blepharomastix ranalis), a round dome, or a longitudinal segment 8.

179 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 10. Herpetogrammatini. A: adult male of Blepharomastix ranalis. B: male genitalia of Eurrhyparodes lygdamis, phallus omitted. C: male genitalia of Herpetogramma licarsisalis, vesica of phallus everted. D: female genitalia of Hileithia cf. obliqualis. E: female geni- talia of H. licarsisalis. F – G: schematic hindlegs, modifed from Lewvanich 1981, Fig. 18. H – I: schematic antennae. — Scale bars: A, G, H – 5 mm; B – E – 500 µm.

The modifed antenna base in males is apomorphic: 62:1, more than one kind of hairpencil chaetae present; the pedicellus is erect, long, with oblong scales emerg- and 99:1, strongly sclerotized colliculum. ing from its distal margin (Hymenia perspectalis (Hüb- Description. The imagines are small (forewing length ner, 1796)) or medially and posteriorly from its base about 9 mm) with dark brown wings contrasted with white (Spoladea recurvalis); the anterior (H. perspectalis) or forewing markings in the median and postmedian lines, medial edge (S. recurvalis) of the pedicellus is raised to and a white transverse band in the hindwing; the head a transverse rim; the basal fagellomeres have a pointy and legs are contrastingly marked. The basal valva costa protrusion on anterior side; and the fagellum is directed is extended into an elongate, ventrad rod that serves as posteriad, giving the antenna a geniculate appearance. dorsal joint with the vinculum. The hairpencils are com- The parsimony trees do not have 40:1, but they add plex, consisting of several sclerotized pads partly with 61:1, two or more hairpencil sclerites on each side of the parallel lines of sclerotized ridges, bearing distinct bun- genitalia, articulated with each other via membranes; dles of long, characteristically bent chaetae; the anterior

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Fig. 11. Hymeniini (B, C, H) and Asciodini (A, D – G). A: adult female of Arthromastix lauralis. B: male of Hymenia perspectalis. C: head of male H. perspectalis. D: head of male Asciodes cf. gordialis. E: male genitalia of As. cf. gordialis, phallus omitted. F: 8th abdominal segment of As. cf. gordialis. G: female genitalia of Ar. pactolalis. H: female genitalia of Spoladea recurvalis. — Scale bars: A, B – 5 mm; E – H – 500 µm.

181 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

half of corpus bursae has a short (S. recurvalis) or long of many genera, the postmedial line is roundly concave (H. perspectalis) transverse ridged signum, the posterior where it crosses the anal fold, rather than angulate. The half of the corpus bursae granulose (H. perspectalis) or sacci tympani are hemispherical and clearly defned; they densely studded with needle-like spikes (S. recurvalis); are smaller and deeper than in most Spilomelinae. The the antrum has a longitudinal, non-sclerotized strip. uncus head is bicapitate or has two separate heads (with a Hymeniini exhibits a typical wing pattern with a dark single head and a central dorsoventral, chaetae-free strip to light brown ground colour interrupted by a broad white in Arthromastix lauralis (Walker, 1859) and Ceratocilia postmedial line on fore- and hindwing; abdomen dorsally sixolalis (Schaus, 1912)); the costa base and vinculum brown with a white band on the segments’ posterior mar- saccus are infated; the transtilla is large and circular or gin. strap-like, with a broad median connection (slim in C. six­ Systematics. Based on our phylogenetic analyses, we olalis). The fbula in the center of the valva is connected place Hymenia Hübner, 1825 (3 spp.) and Spoladea to the sacculus by a distinct “arch” bowing transversely Guenée, 1854 (2 spp.) in Hymeniini. Hymenia and Spo­ across the valva. This arch may bear from one to three ladea represent a part of Munroe’s (1995) polyphyletic digitate processes, or none (). Signa are usually ei- Hymenia genus group. ther absent (most genera) or present as one arcuate line Food plants. The spotted beet webworm , Hymenia (, Sathria, Bicilia). The ostium bursae and ductus perspectalis (Hübner, 1796), and the Hawaiian beet web- bursae are variously sclerotized. Minimally, the collicu- worm moth, Spoladea recurvalis, are polyphagous, their lum is smooth, elongate, and entire (not ventrally mem- larvae feeding on a large variety of food plants, among branous). In some genera, the ostium is fanked by two them several important crops like Amaranthus, Beta, So­ plates or entirely surrounded by wrinkled sclerites, and lanum tuberosum, Spinacia, Xanthosoma and Zea mays the colliculum may be fused with more extensive scleroti- (Solis 2006, 2008). zation along the ductus bursae, which is always shorter Remarks. The extension of the basal valva costa into than the corpus bursae. The posterior end of the corpus an elongate, ventrad rod, serving as dorsal joint with the bursae is often sclerotized with granules or spinules. vinculum, is also present in most investigated Margaro- Systematics. Based on our phylogenetic results we place niini, in Arthromastix lauralis (Asciodini), Samea ec­ Arthromastix Warren, 1890 (2 ssp.), Asciodes Guenée, clesialis Guenée, 1854 (Nomophilini), and Prophantis 1854 (5 spp.) and Arthromastix pactolalis (Guenée, 1854) xanthomeralis (Trichaeini). comb.n. here. Furthermore, based on common morpho- The imagines of Hymenia and Spoladea are very sim- logical features, we assign Beebea Schaus, 1923 (1 sp.), ilar externally, and the generic names have been used in- Bicilia Amsel, 1956 (4 spp.), Ceratocilia Amsel, 1956 terchangeably in the literature. Despite these superfcial (8 spp.), Ceratoclasis Lederer, 1863 (9 spp.), similarities between the adults of Spoladea and Hymenia, Hampson, 1899 (1 sp.), Laniipriva Munroe, 1976 (1 sp.), their genitalia are signifcantly different, and the two gen- Loxomorpha Amsel, 1956 (4 spp.), Maracayia Amsel, era should be kept separate. 1956 (2 spp.), Psara Snellen, 1875 (36 spp.) and Sathria Lederer, 1863 (3 spp.) to Asciodini. We transfer fve genera from Munroe’s (1995) Siga 4.2.15. Asciodini Mally, Hayden, Neinhuis, group: Beebea, Laniifera, Laniipriva, Loxomorpha and Jordal & Nuss trib.n. Maracayia. Males of all the species have the fbula con- nected to the sacculus by an arch, valvae oval in shape Type genus: Asciodes Guenée, 1854 or with a basally infated costa, enlarged transtilla bases (exept Loxomorpha), and the forewing PM line rounded Synapomorphies. 5:1, sensillar setae on basal antenno- basad on the anal fold. Females have a sclerotized la- meres of male > 50% relative to diameter of basal anten- mella postvaginalis, except in Laniipriva. The known nomeres; 72:1, extension (process in some cases) of dor- larvae feed on Cactaceae (Caryophyllales) as borers or sodistal sacculus present; 74:1, basal costa infated; 86:0, webworms. The robust form of the genitalia obscures a phallus apodeme sclerotisation reduced to a ventral, lon- key morphological character – the sacculus-fbula arch – gitudinally sclerotized strip; 107:0, corpus bursae with a but it is visible in careful dissection. Unlike most Ascio- granulose sclerotised area. Slow optimization only: 52:1, dini, the hairpencils are either very simple tufts of hairs connection point of transtillum arms broad; 73:2, fbula or absent, and male antennae are not modifed. Like in and dorsodistal sacculus fused. Margaroniini and Spilomelini, the large size of imagines The parsimony trees do not have 107:0, but they add and “robust” genitalia are syndromatic of the internally fve synapomorphies: 8:0, dorsal direction of 3rd labial feeding larval habit. Laniipriva is problematic because palpo­ mere;­ 33:1, male tergite 8 with heterogenous, dis- the female genitalia illustrated by Munroe (1976b: fg. tinct sclerotization pattern; 51:1, transtillum arms rounded; 21) have an unarmed ostium and a bulged colliculum like 61:1, two or more hairpencil sclerites on each side of the in the Siga group, but the female maculation (ibid. fg. 6) genitalia, articulated with each other via membranes; and is typical of Asciodini. 62:1, more than one kind of hairpencil chaetae present. Ceratocilia (considering C. sixolalis) may have a ba- Description. Males of some genera exhibit modifed an- sal position in this tribe, with its simple transtilla and hair­ tennomeres halfway along the fagellum. In the forewing pencils.

182 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Food plants. Asciodini larvae commonly feed on her- Description. The valvae are weakly sclerotized, lens- baceous Caryophyllales. Asciodes gordialis Guenée, shaped, often with futing on ventral half of valva (cf. 1854 feeds mainly on Nyctaginaceae (Bougainvillea Odontiinae); the fbula is strongly sclerotized (Trichaea) Mirabilis, Pisonia), as does Ceratocilia sixolalis (Neea, to weak, bearing simple hairs (absent in some Prophantis Pisonia); two undetermined Ceratocilia species were spp.); the sacculus is scaly; a ridge-like protrusion is run- reared from Rubiaceae, though (Kimball 1965; Janzen ning from near the sacculus base to the valva centre. The & Hallwachs 2009; Robinson et al. 2010). Arthromas­ corpus bursae has a slim longitudinal signum, its anterior tix lauralis feeds on Trichostigma octandrum (Phytolac- end split into two anterolateral legs in some Prophantis caceae) (Bendicho-Lopez 1998). Bicilia is recorded from species; the ductus bursae is broad, narrowing at the pos- Petiveria and Rivina (Petiveriaceae) (Bendicho-Lopez terior end. 1998; Janzen & Hallwachs 2009). Psara feeds on Ama- Systematics. Based on our phylogenetic analyses, we ranthaceae, Nyctaginaceae, Nelumbonaceae, Phytolac- place Prophantis Warren, 1896 (8 spp.) and Trichaea caceae, P. obscuralis also on Convolvulaceae (Janzen Herrich-Schäffer, 1866 (11 spp.) in Trichaeini. & Hallwachs 2009; Robinson et al. 2010). Laniifera Munroe (1967) points to the distinctness of Thlipto­ cyclades (Druce, 1895), Beebea guglielmi Schaus, 1923, ceras and Prophantis, with several misplaced species at- Loxomorpha, and Maracayia species feed on Cactaceae, tributable to Prophantis. Our phylogenetic results reveal especially , with records of Maracayia on other that the African T. xanthomeralis is one of these cases. Caryophyllales (Mann 1969; Janzen & Hallwachs 2009; Here, we remove this and another African species from Lara-Villalón et al. 2016). Thliptoceras and transfer them to Prophantis: Prophantis Remarks. The upturned palpi (8:0) and heterogenous xanthomeralis (Hampson, 1918) comb.n., and Prophant­ male tergite 8 (33:1) are shared with some other taxa (see is coenostolalis (Hampson, 1899) comb.n. Furthermore, diagnoses of Agroterini and Margaroniini). An extension the African Prophantis longicornalis (Mabille, 1900) of the sacculus (72:1) is shared with Eurrhyparodes, but comb.n. is transferred from Syngamia Guenée, 1854. in that genus, it is a free process, not fused with the fbula. ‘Thliptoceras’ fenestratum Aurivillius, 1910 is also mis- This group of uncolorful moths, as circumscribed placed and belongs to one of the non-euspilomeline clad- here, is a Neotropical radiation on Caryophyllales. Mun- es, probably Udeini. roe’s (1995) association of the gracile external feeders is In the parsimony analysis, Desmia falls in Trichaeini, one of his more perceptive groupings; we doubt that he but it is not supported by any unambiguous morphologi- had knowledge of the host records available to us now. cal characters. Munroe probably associated the large-bodied Beebea and A few Neotropical taxa with mimetic maculation and Laniifera with Siga on overall habitus, and he left Loxo­ Rubiaceae-feeding larvae should be investigated as pos- morpha and Maracayia unplaced. sible members of Trichaeini, but we leave them incertae Hymeniini species, although polyphagous, prefer sedis because the morphological evidence is weak and Amaranthaceae and Chenopodiaceae. Therefore, the sis- we did not sequence them. They include Erilusa Walker, ter-group relationship of Asciodini and Hymeniini under some results (Bayesian and implied-weights parsimony 1866 and species misplaced elsewhere, such as Phostria under k = 9 – 13) suggests that feeding on Caryophyllales delilalis (Walker, 1859) and Pilocrocis xanthozonalis is a synapomorphy of the two tribes. Hampson, 1912. Females of Erilusa and P. xanthozona­ lis have a large, complete colliculum, a short, granulose ductus bursae, and two small, round signa, characters that 4.2.16. Trichaeini + (Steniini + Nomophilini) relate them to Prophantis. In Erilusa, the uncus is uni- capitate and the elliptic, dentate fbula is not connected to Synapomorphies. 109:5 (unique), anterior-most signum the sacculus, which exclude it from Asciodini. The uncus a transverse, smooth or dentate arch, with or without cen- varies from capitate in Erilusa to reduced and triangu- tral posteriad leg (if present, then signum Y-shaped) (not lar in P. xanthozonalis, but the uncus is likewise variably found with parsimony). Slow optimization only: 33:1, reduced in Trichaeini. Sacculosia Amsel, 1956 (1 sp.) sclerotization of male tergite 8 heterogeneous (in parsi- shares a fbula and futed valva similar to Trichaea, but mony trees); 67:2, fbula directed towards distal valva; more information is needed in order to investigate this 78:2, general shape of post-basal costa (not the entire hypothetical relationship. dorsal valva edge) convex (not found with parsimony); Food plants. (Butler, 1875), P. 104:0, sclerotisation in ductus bursae absent. octoguttalis (C. Felder, R. Felder & Rogenhofer, 1875) and P. longicornalis are recorded as pests on Coffea ara­ bica (Rubiaceae) and referred to as ‘[coffee] berry moths’; 4.2.17. Trichaeini Mally, Hayden, Neinhuis, alternative hosts are Tricalysia and Bertiera zaluzania, Jordal & Nuss trib.n. Ixora coccinea, Gardenia (Rubiaceae), Duranta plumieri Type genus: Trichaea Herrich-Schäffer, 1866 (Verbenaceae) and Triclisia (Menispermaceae) (Waller et al. 2007; Guillermet 2009). Hinckley (1964) reports Synapomorphies. 70:1, raised ridge running from basal an undescribed Prophantis from boring in Gardenia to dorsodistal sacculus present. fowers and shoots.

183 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 12. Trichaeini (A, D) and Steniini (B, C, E – G). A: adult male of Trichaea sp. B: adult male of Dolicharthria aetnealis. C: male genitalia of Do. punctalis, phallus omitted. D: male genitalia of T. pilicornis. E: male genitalia of Duponchelia fovealis, phallus omitted. F: female genitalia of Du. fovealis. G: abdomen of male Penestola bufalis. — Scale bars: A, B – 5 mm; C – F – 500 µm.

Janzen & Hallwachs (2009) report Trichaea larvae distal sacculus (or its extension) distant from each other, from Psychotria spp., Morinda panamensis and Marga­ non-overlapping. The parsimony trees imply the same, ritopsis microdon (Rubiaceae), with a single record on with both characters unambiguous. each Urticaceae and Celastraceae. Feeding on Rubiaceae Description. Imagines often have long legs; males have is shared with Nomophilini (see also remarks there). a slender, long abdomen. The uncus is single or bicapitate Remarks. The slim longitudinal signum of some Pro­ (Loxostegopsis, Tatobotys) or entirely split (Metasia) and phantis species with its anterior end split into two ante- has bifd chaetae. The valva costa is concave or straight, rolateral legs is paralleled in Syngamia (Nomophilini). in some taxa weakly convex, and the valva is simple with usually zero or one fbula originating from base of valva, or in the Duponchelia group (sensu Hayden 2011) with 4.2.18. Steniini Guenée, 1854 stat.rev. two or three small fbulae at the base of the valva; the Type genus: Guenée, [1845] = Dolicharthria Stephens, 1834 phallus has a caecum. The signum is absent, except in , , and Perisyntrocha, where it is a Synapomorphies. 106:0, sclerotisation in corpus bursae toothed arc. The ostium and ductus bursae lack any other absent. Slow optimization only: 73:0, fbula and dorso- sclerotization.

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In the Duponchelia group, Rs1 is stalked with Rs2+3 fallen plant matter (e.g. Hinckley 1964). Duponchelia in the forewing, and a fovea in the male forewing at the fovealis Zeller, 1847 as a pest is spread through the plant distal end of the discal cell is a recurrent character in sev- nursery trade as it feeds on organic potting soil and suc- eral but not all species; a pair of elongate hairpencils is culent stems and foliage. present dorsal of the vinculum; the ductus bursae is very short; the larvae have the mesothoracic SD2 seta fne and hairlike. 4.2.19. Nomophilini Kuznetzov & Stekolnikov, Systematics. Based on our phylogenetic results, we 1979 stat.rev. place Anageshna Munroe, 1956 (1 sp.), Dolicharthria Type genus: Nomophila Hübner, 1825 Stephens, 1834 (24 spp.), Duponchelia Zeller, 1847 (5 spp.) and Metasia Guenée, 1854 (88 spp.) in Steniini. Synapomorphies. No unambiguous synapomorphies Furthermore, based on morphology we place Apogeshna could be found for this tribe as circumscribed in the Munroe, 1956 (3 spp.), Bradina Lederer, 1863 (87 spp.), Bayesian results. Slow optimization only: 74:1, basal Epherema Snellen, 1892 stat.rev. (1 sp.), Hymenoptychis costa infated. Two sister clades are present in Nomophil- Zeller, 1852 (4 spp.), Loxostegopsis Dyar, 1917 (6 spp.), ini, Syngamia + (Ategumia + (Bocchoris + (Diasemia Penestola Möschler, 1890 (3 spp.), Steniodes Snellen, + Diasemiopsis))) and Desmia + ((Mecyna + Arnia) + 1875 (9 spp.), Symmoracma Meyrick, 1894 (1 sp.) and (Samea + Nomophila)). In the former clade, no synapo- Tatobotys Butler, 1881 (11 spp.) here. Bradina is unusual morphies or characters from slow optimization are found, in possessing a signum, which could be plesiomorphic. but the latter clade is characterised by the synapomor- The genera Duponchelia, Hymenoptychis, Penestola and phies 67:0, fbula ventrally directed towards sacculus or Tatobotys are considered to be closely related: they have distal sacculus, and 95:0, longitudinal membranous strip two or three small fbulae, a pair of narrow vincular andro- in the antrum sclerotisation absent. conia, forewing Rs1 stalked with Rs2+3 and frequently a Nomophilini as circumscribed here is not monophyl- fovea, and the larvae are semiaquatic in swamps. We did etic in the parsimony trees. not examine the type species of Nacoleia, N. rhoeoalis A core Nomophilini s.str. consisting of Mecyna, No­ (Walker, 1859), but certain important species in this large mophila, and Samea (without Desmia or Syngamia) has genus (84 spp.) such as N. octasema (Meyrick, 1886) and several synapomorphies in both the Bayesian and parsi- N. charesalis (Walker, 1859) belong to Steniini based on mony trees: 5:1, sensilla of male antennae elongate; 89:1, the position of the fbula, absence of a signum, and sap- vesica with multiple cornuti; 100:0, colliculum evenly rophagous larval habits. sclerotized all around, without membranous strip; and Lederer, 1863 (93 spp.) probably belongs 109:1, signum longitudinal and granular. to Steniini. We have not studied the type species P. viola­ Description. Small to medium-sized moths. The wing lis Lederer, 1863, but P. signiferalis (Wallengren, 1860) pattern is reticulated in many Neotropical genera. The as illustrated by Clarke (1986: fgs. 56, 57) shares the Diasemia group have sacci tympani normally developed maculation, a broad and deep saccus, ornate valvae, com- (Bocchoris), small (Ategumia) or absent (Diasemia, plex hairpencils, and a corpus bursae with spicules but Diasemiopsis), with the fornix tympani in contact with no single signum. It is related to a group of Steniini that the tympanic frame all around. The male genitalia have have a broad saccus and ornate valvae that includes cer- a conical to capitate uncus (reduced in Ategumia, Dia­ tain Steniodes species (S. mendica (Hedemann, 1894), S. semia and Bocchoris), uni- to bicapitate, uncus head na- acuminalis (Dyar, 1914)), Camptomastix Warren, 1892 ked or with simple and/or bifurcate chaetae; the valvae and Symmoracma Meyrick, 1894. Lipararchis Meyrick, are ovate, mostly with a convex costa; the fbula is well- 1934 (2 spp.) might belong here too. developed, straight to arched and emerging from near Food plants. Little is known about the feeding habits of the costa base (small in Desmia, absent in Bocchoris, Steniini. Dolicharthria punctalis (Denis & Schiffermül- Diasemia and Diasemiopsis). In the female genitalia, the ler, 1775) preferably on wilting leaves of different plants corpus bursae has a granulose central area or an elon- (Hasenfuss 1960). Metasia corsicalis (Duponchel, 1833) gate signum, longitudinal or transverse in orientation, is reported to feed on detritus (Leraut 2012). Nacoleia in Diasemia and Bocchoris invaginated to form a spine; charesalis feeds on rotting leaves and bores in turmeric the colliculum is sclerotized, in Nomophila and ‘Samea’ stems, and N. octasema consumes inforescences of ba- multiplicalis (Guenée, 1854) with an apomorphic blind nanas (Paine 1964; Hiremath et al. 1990; Kumar et al. anterolaterad evagination (diverticulum sensu Munroe 1996; Tominaga 2002). The absence of records by itself 1973); the antrum is strongly sclerotized, broad tubular suggests that the saprophagous habit is common, because or barrel-shaped. such larvae would be easy to overlook. One group is par- Systematics. Based on our phylogenetic results we place ticularly interesting: the larvae of the Duponchelia group Arnia Guenée, 1849 (1 sp.), Ategumia Amsel, 1956 (10 (sensu Hayden 2011) are detritivores in marshes and spp.), Bocchoris Moore, 1885 (31 spp.), Desmia West- intertidal environments of mangrove swamps (Murphy wood, 1832 (89 spp.), Diasemia Hübner, 1825 (13 spp.), 1990). They are often associated with the Avicennia zone Diasemiopsis Munroe, 1957 (2 spp.), Mecyna Double- of mangrove forests, which is inundated at high tides. The day, 1849 (34 spp.), Nomophila Hübner, 1825 (14 spp.), larvae live on the ground and feed on rich soil and juicy Samea Guenée, 1854 (28 spp.) and Syngamia (25 spp.)

185 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 13. Nomophilini. A: male genitalia of Desmia tages, phallus omitted. B: male genitalia of Ategumia ebulealis, phallus omitted. C: ab- domen segments 4 – 7 of male Samea ecclesialis. D: female genitalia of Desmia sp. E: female genitalia of Nomophila noctuella. F: phallus of Mecyna lutealis. — Scale bars: 500 µm. here. Furthermore, we place Crocidocnemis Warren, spp.), Pardomima Warren, 1890 (16 spp.), Perisyntrocha 1889 (2 spp.), Warren, 1892 (10 spp.), Diasemi­ Meyrick, 1894 (4 spp.), Pessocosma Meyrick, 1884 (4 odes Munroe, 1957 (4 spp.), Diathrausta Lederer, 1863 spp.) and Snellen, 1880 (15 spp.) in Nomo- (20 spp.), Hübner, 1825 (14 spp.), Mimophobe­ philini based on morphological characters. tron Munroe, 1950 (1 sp.), Warren, 1890 (15 Arnia Guenée, 1849 was synonymized with Stenia spp.), Warren, 1892 (1 sp.), Nothomastix Duponchel, 1845 (a synonym of Dolicharthria Stephens, Warren, 1890 (5 spp.), Parapilocrocis Munroe, 1967 (3 1834) by Rebel (1901), a decision that was revoked by

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Amsel (1952). We fnd Arnia as sister to Mecyna, and less commonly) a transverse “moustache” with two more they share a number of morphological features, such as or less connected sections, found in Samea castellalis shape of uncus and fbula, multiple dentiform cornuti in Guenée, 1854 and (Zeller, 1852). the phallus, a central granulose area in the corpus bursae, An infated basal costa (74:1) is shared with some Ste- and a short broad, sclerotized ductus bursae. niini (Dolicharthria and Metasia). Food plants. Larvae are leaf-rollers mainly on Fabaceae, Samea is paraphyletic with respect to Nomophila in Onagraceae, Rubiaceae and Vitaceae, with occasional our phylogenetic results. Both S. multiplicalis and most records from Begoniaceae, Cordiaceae and Malvaceae species of Nomophila (see Munroe 1973) have a large, (Kimball 1965; Allyson 1984; Solis 2008; Janzen & deeply arched fbula, uncus without large bifd chaetae, Hallwachs 2009; Hayden 2014). Ategumia feeds mainly and the colliculum extended into a diverticulum. on , furthermore on Rubiaceae and Big- Nomophila was revised by Munroe (1973), the Af- noniaceae, with single records on Fabaceae, Piperaceae rican Pardomima species by Martin (1955), Syngamia and Urticaceae (Janzen & Hallwachs 2009). Boccho­ forella (Stoll in Cramer & Stoll, 1781) and its variations ris inspersalis (Zeller, 1852) is reported from Malva- by Heppner (2010). This tribe generally corresponds to ceae, Fabaceae and Amaranthaceae (Ghesquière 1942; the Samea group of Munroe (1995). Wagner et al. 2008; Robinson et al. 2010). Diasemia is A simple tubular colliculum and transverse sig- recorded from Asteraceae, Plantaginaceae and Lecythi- num (similar to that in S. castellalis) are also found in daceae (Ghesquière 1942; Robinson et al. 2010), Synga­ Diasemiodes, Diathrausta, and Perisyntrocha (Munroe mia mainly on Rubiaceae, with further records on Acan- 1956), so Nomophilini seems to be a better tribe for these thaceae and Asteraceae (Janzen & Hallwachs 2009), genera than Steniini. On the other hand, the absence of Pardomima was recorded from coffee (Rubiaceae) (Mar- checkered maculation suggests that further investigation tin 1955). Larvae of the monotypic Mimophobetron feed is needed. on different species of Rubiaceae (Janzen & Hallwachs 2009). Nothomastix klossi is recorded from Psychotria (Rubiaceae) (Miller et al. 2007). 4.2.20. Munroe’s (1995) group ‘Samea’ multiplicalis (Guenée, 1854) and Niphograp­ ta albiguttalis (Warren, 1889), whose larvae are used in Apart from Syllepte (see below), only the core of Mun- biological control of aquatic weeds, may represent an roe’s (1995) Eulepte group is not represented in our mo- aquatic lineage, related to Crocidocnemis, whose larvae lecular sampling (three other genera are transferred to are not known. Furthermore, Hydririni). For this reason, we do not formally propose was reported to feed on leaves of the aquatic fern Azolla it as a tribe. The genera Eulepte Hübner, 1825 (6 spp.), fliculoides (Salviniaceae) (Farahpour-Haghani et al. Praeacrospila Amsel, 1956 (4 spp.), Leucochromodes 2016). Amsel, 1956 (8 spp.), and Mesocondyla Lederer, 1863 In the clade Trichaeini + (Nomophilini + Steniini), (2 spp.) have oval to moderately attenuate valvae, uncus the larvae of most Trichaeini and of several Nomophil- with bifd chaetae, and a pair of simple vincular andro- ini (especially the early-diverging Desmia and Synga­ conia with long, hairlike setae. The saccus is elongate in mia) feed on Rubiaceae. Considering this relationship, Mesocondyla and Eulepte, and perhaps most distinctive- we hypothesize that this is the primitive host family for ly, most taxa (except M. dardusalis) have two inwardly Nomophilini or maybe for the entire clade Trichaeini + curved fbulae closely set together: one an extension of (Nomophilini + Steniini), and that Nomophilini has radi- the sacculus, the other from the face of the valva, just in- ated onto other hosts. However, larvae of Steniini, as far side and curving in parallel with the saccular fbula. The as known, are detritivorous. ductus bursae is elongate in Eulepte and Mesocondyla, Remarks. The common form of the male genitalia of short in Leucochromodes and Praeacrospila, and the sig- Nomophilini is fairly nondescript, with few striking num is absent or double. The maculation is yellow with characters: the uncus is usually simple or weakly bifd, a darker postmedial area, which however also occurs the valvae are elliptical, and there is one curved fbula or in other taxa (e.g. Lygropia species). Zenamorpha dis­ none. However, particular genera or genus groups show cophoralis (Hampson, 1899) is another possible member interesting characters, such as modifcation or loss of the of this group, considering the male genitalia, although it uncus (Nomophila, Ategumia), distally concave valva could also belong to Trichaeini. (Sa­mea cancellalis, Diacme), or one pair of apical cor- nuti in several genera (e.g. Mecyna, Mimorista, Samea, Epi­pagis). Likewise, the shape of the signum may be in- 4.2.21. Syllepte Hübner, 1823 formative above the genus level. The maculation of the “core” Nomophilini is characteristically chequered, but Syllepte, the type genus of “Sylleptinae”, is a large poly- other taxa (Desmia, Syngamia, Mimophobetron) show phyletic genus within Spilomelinae, containing 199 valid other patterns. The “core” Nomophilini is well-charac- species (Nuss et al. 2003 – 2019). The identity of the ge- terized by female genitalia. The colliculum is tubular nus is ambiguous as the type material of its type species, with one or two lateral pockets, and the signa have two Syllepte incomptalis Hübner, 1823 (and not Phalaena forms: commonly a longitudinal granular strip or (much amando Cramer, 1779, as erroneously stated by Kirti

187 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

& Gill 2007), is lost (Groll 2017). The illustrations of pattern; 55:1, juxta split 10 – 60% of its length (only in the male specimens of this species depicted in Hübner the parsimony trees); 60:1, partly sclerotized chaetose (1819 – 1823: 18, pl. [50] fgs. 285, 286) are diffcult to hairpencils articulating with the anterior edge of the vin- associate to any known species. The maculation resem- culum tegumen connection present; 99:1, strongly scle- bles, to some degree, Bocchoropsis Amsel, 1956 and rotized colliculum anterior of the antrum and posterior of specimens of the Polygrammodes eleuata (Fabricius, the attachment of the ductus seminalis present (not found 1777) species group. with parsimony). Slow optimization only: 8:1, direction of third labial palpomere porrect (not found with parsi- mony); 32:1, anterior edge of male tergite 8 deeply emar- 4.2.22. Genera removed from Spilomelinae ginate (only in the parsimony trees); 109:6, anterior-most signum broad, medially constricted, resembling puck- Aporocosmus Butler, 1886 is transferred to Odontiinae, ered lips (unique; not found with parsimony). where it is related to Thesaurica Turner, 1915. Orthora­ Description. A retinacular hook (frenulum hook sensu phis Hampson, 1896 is transferred to Lathrotelinae. Forbes 1926) is present in the male forewing of 13 of the Hydropionea Hampson, 1917, Plantegumia Am- 18 investigated Pyraustinae. The mesothoracic tibia in sel, 1956 and Munroe’s (1995) “undescribed genus ex males has a hidden hairpencil (Ohno 2000; Frolov et al. Boeotarcha Meyrick” appear to form a group of aber- 2007). The fornix tympani is recessed within tympanic rant Glaphyriinae. Munroe (1995) probably placed them frame. The hemispherical sacci tympani tend to be large in Spilomelinae because the males have the gnathos re- and deep, especially in Pyraustini and Portentomorphini. duced to a transverse band or absent. However, the males The shape and large size is paralleled in some Odontiinae. possess a retinacular hook, but they do not have the syna- The degree at which the praecinctorium of the tympanal pomorphies of Pyraustinae. We transfer them to Glaphy- organ is bilobed is neither distinctive for Spilomelinae riinae s.l. (Regier et al. 2012). This is supported by 1) nor for Pyraustinae, so that this character is unreliable for narrow valvae with apically separate costa and sacculus, distinguishing the two subfamilies. Male genitalia have and 2) tympanal organs with large, mesal sacci tympani a transtilla inferior (sensu Marion 1954; absent from Te­ and large puteoli. A species of Hydropionea has been tridia and many other taxa) and a sella (sensu Marion raised on Capparis unifora (Janzen & Hallwachs 2009), 1952) on the inner surface of the valva, often with strong which fts with Glaphyriinae s.l., a clade best defned as piliform or spatulate hairs (editum sensu Marion 1952); a radiation on mustard-oil producing (Regier the editum is absent from many taxa. The female genita- et al. 2012). lia have a long, coiled ductus bursae (absent in several Phaedropsis leialis (Dognin, 1906) and Lygropia taxa, e.g. , Ostrinia, Uresiphita). Deciduous cor- murinalis Schaus, 1912 are related and misplaced in Spi- nuti are present. lomelinae. They have a male retinacular hook, tympanal Appendix bursae present, emerging from the anterior organs with the fornix at the same level as the venula pri- ductus bursae (and not from the corpus bursae) in Te­ ma, gnathos with medial process, unmodifed valvae, and tridia, Euclastini and Portentomorphini, or laterally from an ediacaroid signum. This combination of characters is the corpus bursae in Pyraustini and Uresiphita. Signum very puzzling; we tentatively place them in Pyraustinae broad rhombical (Pyraustini), ‘puckered lips’-shaped in incertae sedis. The host (Gouania Jacq.: Rhamnaceae; Euclastini and Tetridia Warren, 1890, or ediacaroid in Janzen & Hallwachs 2009) is not informative. Portentomorphini and Uresiphita. Certain species belong to Pyraustinae incertae se- The shapes of the teguminal ridges in Solis & Maes dis. Lygropia fusalis Hampson, 1904 and related species (2003: character 9) seem to be good for diagnosing tribes. are Pyraustinae, based on the editum of comb-tipped Remarks. Plesiomorphic characters shared with the non- scales on the sella, concave costa, the deeply invaginated euspilomeline clades in Spilomelinae are: absence of a sacci tympani, and data from the nuclear EF-1a gene. sclerotized strip on the pleural membranes of segment 8 Blepharomastix haedulalis (Hulst, 1886) is another with (present in Euclastini); costa straight to concave; saccus typically pyraustine male genitalia. Females of both taxa of vinculum broadest at the base, without a basal con- have no signum, so their placement in Pyraustinae was striction. Deciduous cornuti are paralleled in the Spilo- overlooked. melinae ‘Syllepte’ adductalis (Walker, 1859) and Pyc­ narmon pantherata (Agroterini). The ediacaroid signum of Portentomorphini and Uresiphita is shared with the non-euspilomeline Spilomelinae. 4.3. Pyraustinae Meyrick, 1890

Type genus: Pyrausta Schrank, 1802 4.3.1. Tetridia Warren, 1890

Autapomorphies. 29:1, large, oval pleural scale tufts Synapomorphies. 22:1, fornix tympani surface recessed on each side of the male abdominal segment 7 present, within the frame (unique); 33:1, heterogenous scleroti- with an opening in its anterior centre (unique); 57:1, ba- zation of male tergite 8, i.e. with distinct sclerotization sal saccus constricted; 58:1, ratio between saccus length

188 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Fig. 14. Tetridia (A, C, D, F, H) and Euclastini (B, E, G). A: adult male of Tetridia vinacealis. B: adult female of Euclasta gigantalis. C: male genitalia of T. vinacealis. D: phallus of T. vinacealis. E: male genitalia of E. splendidalis. F: 7th abdominal segment of male T. vi­na­cealis. G: female genitalia of E. splendidalis. H: female genitalia of T. vinacealis. — Scale bars: A, B – 5 mm; C – H – 500 µm.

189 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

and sacculus breadth > 1; 63:1, presence of a pair of body and wings fuscous; legs ferruginous; both wings sclerotized, hair-studded hairpencils articulating with with a series of terminal black spots; ante- and postme- the anteromedian edge of the saccus; 66:0, general shape dian lines on the forewing distinctly different in the dor- of fbula broadly triangular; 72:1, extension (process in sal half. Kirti & Gill (2007), on the other hand, state that some cases) of dorsodistal sacculus present; 76:1, long, “this species [T. caletoralis] drastically differs from the sometimes loosely arranged chaetae on surface of costal type species of the genus Tetridia Warren i.e., vinacealis base present; 86:1, phallus apodeme sclerotisation re- Moore” (Kirti & Gill 2007: p. 266). Many adult Pyraus- duced to a ventral, longitudinally sclerotized strip; 87:1, tinae and Spilomelinae exhibit a wing pattern similar to a distinct sclerite in the posterior phallus apodeme pre- that of T. vinacealis, and we assume that Kirti & Gill sent; 89:0, vesica with single cornutus; 92:1, ventral end (2007) misidentifed their material. A revision of the ge- of papillae anales larger than dorsal end; 100:0, longitu- nus might bring certainty on this matter and might an- dinal membranous strip in the colliculum sclerotisation swer the question whether this taxon should be placed in absent; 108:1, two or more signa; 110:1, second signum a separate tribe. (located posterior of frst signum) slim, strip-like. Slow Food plants. Tetridia caletoralis is recorded from Sho­ optimization only: 25:0, venulae secundae convergent; rea robusta () (Robinson et al. 2010). 49:1, uncus attached to tegumen as a broad, smooth tran- Remarks. The paired hairpencil-like structures attached sition; 90:2, orientation of everted papillae anales poste- anteromedially to the vinculum are a plesiomorphy riad; 112:1, appendix bursae present on anterior ductus shared with Lamprosema in Spilomelinae: Hydririni; the bursae. ‘puckered lips’-shaped signum is shared with Euclastini; Description. The antennae are longer than the forewing. the attachment of the appendix bursae to the anterior end The uncus has spatulate chaetae in addition to simple, of the ductus bursae is shared with Euclastini and Porten- hair-like chaetae; the sacculus broad and triangular, oc- tomorphini. cupying the ventral valva base, and the centre of the dor- sal sacculus edge has a robust spine pointing dorsally to- wards the uncus, and a second, more fragile, curved spine 4.3.2. Euclastini Popescu-Gorj & Constanti- further towards the distal sacculus; the ventral vinculum nescu, 1977 stat.rev. anteromedially has paired hairpencil-like structures. The signum is broad, medially constricted, resembling puck- Type genus: Euclasta Lederer, 1855 ered lips (‘spectacles-shaped’ in Popescu-Gorj & Con- stantinescu 1977); the appendix bursae emerges at the Synapomorphies. 10:1, length of maxillary palpi minute anterior end of the ductus bursae. The tegumen ridges to obsolete, cannot hypothetically come in contact with cannot be discerned, because the scale-bearing lateral each other (not found with parsimony); 16:0, metatibial felds of the tegumen are expanded and compress the me- proximal inner spur shorter than half of tibial segment sal area into a narrow strip. Therefore, the ridges could be between this and the distal spur pair; 47:2, multifurcate either absent or fused. structure of uncus head chaetae (unique); 65:0, absence Systematics. So far, we place only two species of Te­ of fbula emerging from dorsal valva base near costa base tridia here, the type species T. vinacealis (Moore, 1877), (not found with parsimony); 78:2, general shape of post- and T. caletoralis (Walker, 1859). basal costa (not the entire dorsal valva edge) convex. The identity of T. caletoralis is still not fully clear, Slow optimization only: 18:1, scale brush at costal base as the type material at NHMUK could not be traced. We of forewing underside in males formed into a retinacular used the DNA extract of voucher specimen WPH209 from hook (not found with parsimony); 44:1, shape of uncus Haines & Rubinoff (2012) for our molecular dataset, but capitate (not found with parsimony); 49:1, attachment of specimens available for morphological study could not uncus to tegumen broad, smooth transition; 109:6, shape be confrmed as conspecifc with specimen WPH209, of anterior-most signum broad and medially constricted which had no genitalia left for investigation. Instead, for (only in the parsimony trees); 112:1, appendix bursae the morphological investigation we used DNA-barcoded present on anterior ductus bursae. material that was in the nearest neighbour BIN of speci- Description. The imagines are long-legged and gracile men WPH209 in the Barcode of Life Database (BOLD). with narrow, apically rounded forewings with brown kirti & Gill (2007) transferred T. caletoralis to Pata­ dorsal ground colour, traversed by a whitish band from nia, based on material from the Natural History Museum wing base to apex, and two dark discal spots. The fore- London. We have seen the NHMUK Pyralidae slide no. wings are held parallel to the frontally raised body when 19900 (male) to which Kirti & Gill (2007) likely refer resting, somewhat resembling Lineodes (Spilomelinae: and agree that this taxon belongs to Patania or at least to Lineodini). The uncus has a bulbous head with multifd Agroterini. However, we have doubts about the correct chaetae; the valvae are trapezoid, with the straight ven- identifcation of the material referred to by Kirti & Gill tral valva edge parallel to the straight costa that spans the (2007), and we have not seen the specimen from which basal half of the dorsal valva edge; and the distal dorsal NHMUK Pyralidae slide no. 19900 originates. Shibuya valva edge runs more or less straight towards valva apex. (1928) mentions four characters in which T. vinacealis, The fbula is absent. The signum is broad, medially con- the type species of Tetridia, differs from T. caletoralis: stricted and laterally slimly extended, resembling puck-

190 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

ered lips (‘spectacles-shaped’ in Popescu-Gorj & Con- costa base (only in parsimony trees); Slow optimization stantinescu 1977); the appendix bursae emerges at the only: 109:3, anterior-most signum transverse rhombical anterior end of the ductus bursae. to cross-shaped, with longitudinal axis shorter than or The tegumen ridges are closely parallel, connected by equally long as transverse one. anterior crossbar at junction of V, like a two-legged “Y”. Systematics. Euclasta (17 spp.) is the only included ge- nus. 4.3.4. Portentomorphini + Uresiphita Food plants. All known host plant records are from Apo- cynaceae, with Euclasta splendidalis (Herrich-Schäffer, Synapomorphies. 46:0, chaetae on surface of uncus 1848) on Periploca graeca, E. warreni Distant, 1892 on head(s) absent; 61:1, two or more hairpencil sclerites on Acokanthera oppositifolia and E. maceratalis Lederer, each side of the genitalia (articulated with each other via 1863 on Gymnanthera nitida (Popescu-Gorj & Constan- membranes); 82:1, costa detached from valval area, the tinescu 1977; Common 1990). costa protruding freely dorsad (unique). Slow optimiza- Remarks. The valva shape is somewhat paralleled tion only: 21:1, lobulus on lateral edge of tympanal case in Chilopionea Munroe, 1964 and some species of present; 44:1, shape of uncus capitate; 114:1, posterior Chilochroma Amsel, 1956 (Pyraustini). The ‘puckered point of attachment of appendix bursae on corpus bursae. lips’-shaped signum is shared with Tetridia, and the ori- gin of the appendix bursae at the anterior end of the duc- tus bursae is shared with Portentomorphini and Tetridia. 4.3.5. Portentomorphini Amsel, 1956 stat.rev. Maes (2000) postulates a close relationship between Paschiodes Hampson, 1913, Duzulla Amsel, 1952 and Type genus: Portentomorpha Amsel, 1956 Euclasta. We have not studied Duzulla and the fgure and description in Amsel (1952) are inconclusive, and we Synapomorphies. 5:1, sensillar setae of males at basal therefore refrain from speculation. We have seen material antennomeres > 50% relative to diameter of basal anten- of Paschiodes, and the presence of bifd chaetae (instead nomeres (not found with parsimony); 46:0, chaetae ab- of multifd chaetae as in Euclastini) makes this relation- sent from surface of uncus head(s); 67:3, fbula generally ship unlikely. Only one of the fve species of Paschiodes oriented dorsally towards tegumen or uncus. Slow opti- (P. ugandae Maes, 2005) exhibits an appendix bursae, mization only: 51:3, transtillum arms large rectangular, and this emerges from the side of the corpus bursae, a medioventrally with fnger-like process (transtilla infe- character corresponding to Pyraustini and to Uresiphita rior sensu Marion 1954). The parsimony trees add two (see remarks under Portentomorphini). Until an analysis characters: 21:1 and 82:1. on the phylogenetic relationship of Paschiodes is done, Description. The male genitalia have the costa detached we refrain from placing the genus in one of the proposed from the valva and projecting freely dorsad, bearing a tribes. terminal feld of setae. A thin, elongate, curved, often resembles Euclasta in the bulbous un- articulated fbula emerges from the centre of the dorsal cus head, the valva shape and the absence of a fbula and valva edge, reaching dorsad; the actual valva consists of sella in the male genitalia, and in the appendix bursae the far dorsad reaching sacculus which ends in a termi- emerging from the anterior end of the ductus bursae as nal setose feld in the valva apex; the sacculus is large well as the ‘puckered lips’-shaped signum. The uncus and membranous. The uncus is narrow, naked, and often chaetae are bisetose and not multisetose as in Eucla­ distally forked. In addition to these synapomorphies, the sta. Saucrobotys larvae also feed on Apocynaceae, with appendix bursae emerges at the anterior end of the ductus S. futilalis (Lederer, 1863) on Apocynum and Asclepias bursae close to the transition into the corpus bursae, in syriaca, where the larvae live gregarious in a nest made Pioneabathra J.C. Shaffer & Munroe, 2007 laterally at- from leaves and silk; the plant associations for larvae tached to the corpus bursae. The signum is a four-armed and pupae of S. fumoferalis (Hulst, 1886) are doubtful star in Hyalobathra Meyrick, 1885 and Cryptosara E. L. (Munroe 1976a). Because of the bisetose uncus chaetae, Martin in Marion, 1957, an ediacaroid sclerite in Por­ we refrain from placing Saucrobotys in Euclastini. Eu­ tentomorpha, and in Pioneabathra and Isocentris flalis clasta has been revised by Popescu-Gorj & Constanti- (Guenée, 1854) there are two large, opposing granulose nescu (1977), the two species of Saucrobotys are treated areas. The maculation is basically yellow but often has a in Munroe (1976a). distinctively red or orange postmedial area (or entirely pink: e.g. Hyalobathra unicolor (Warren, 1895)).The tegumen is short and evenly sclerotized, without dorsal 4.3.3. (Portentomorphini + Uresiphita) + ridges. Pyraustini Systematics. Based on our phylogenetic results we place Hyalobathra (21 spp.), Cryptosara (3 spp.) and Porten­ Synapomorphies. 20:1, splitting of praecinctorium tomorpha (1 sp.) in Portentomorphini. Munroe (1976a) weak to absent. 25:1, Course of venulae secundae par- recognizes a group of related genera comprising Porten­ allel or diverging in posterior half (only in parsimony tomorpha, Cryptosara, Isocentris Meyrick, 1887 (7 spp.) trees); 65:1, fbula emerging from dorsal valva base near and Hyalobathra. We concur with Munroe’s (1976a)

191 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Fig. 15. Portentomorphini (A, C, E, G) and Pyraustini (B, D, F). A: adult female of Pioneabathra olesialis. B: adult male of Pagyda sp. C: male genitalia of Hyalobathra illectalis, phallus omitted. D: male genitalia of Achyra nudalis. E: sternites 6 – 8 of male Cryptosara caritalis. F: female genitalia of . G: female genitalia of H. illectalis. — Scale bars: A, B – 5 mm; C – G – 500 µm.

192 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

composition of this group, here defned as Portentomor- longer than the longitudinal axis. The tegumen ridges are phini, and further include the African monotypic genus parallel, widely spaced, and not connected. Pioneabathra. Systematics. According to our phylogenetic results, we Food plants. Most food plant records are from Phyllan­ place the following taxa in Pyraustini: Achyra Guenée, tha­ceae (Malpighiales): The monotypic Portentomor­ ­pha 1849 (19 spp.), Anania (117 spp.), Hyalorista Warren, feeds on Margaritaria nobilis, Hyalobathra species on 1892 (5 spp.), Loxostege (90 spp.), Oenobotys Munroe, Glo­chi­dion and Phyllanthus, but they are also recorded­ 1976 (5 spp.), Ostrinia Hübner, 1825 (21 spp.), Pagyda from Abrus (Fabaceae) and (Asteraceae); Iso­ Walker, 1859 (26 spp.), Paracorsia Marion, 1959 (1 centris flalis (Guenée, 1854) and the monotypic Pionea­ sp.), Psammotis Hübner, 1825 (8 spp.), Pseudopyrausta bathra on Flueggea, the latter also on Solanum (Sutrisno Amsel, 1956 (6 spp.), Pyrausta (341 spp.) and Sitochroa & Horak 2003; Janzen & Hallwachs 2009; Robinson et Hübner, 1825 (10 spp.). al. 2010). Based on morphology, we furthermore place the fol- Remarks. The origin of the appendix bursae at the ante- lowing genera in Pyraustini: Adoxobotys Munroe, 1978 rior end of the ductus bursae is shared with Tetridia and (3 spp.), Aglaops Warren, 1892 (4 spp.), Anamalaia Euclastini. Munroe & Mutuura, 1969 (1 sp.), Arenochroa Munroe, We consistently fnd Uresiphita Hübner, 1825 to be 1976 (1 sp.), Aurorobotys Munroe & Mutuura, 1971 sister to Portentomorphini in our phylogenetic analyses. (2 spp.), Callibotys Munroe & Mutuura, 1969 (3 spp.), Uresiphita does not share any of the unusual synapo- Carminibotys Munroe & Mutuura, 1971 (1 sp.), Ceutho­ morphies of the other Portentomorphini, and we there- botys Munroe, 1978 (1 sp.), Chilochroma Amsel, 1956 fore do not include it in the tribe. Slow optimization in (4 spp.), Chilocorsia Munroe, 1964 (1 sp.), Chilopionea WinClada results in the following three synapomorphies: Munroe, 1964 (1 sp.), Butler, 1879 (19 spp.), 5:1, length of sensillar setae at basal antennomeres rela- Lederer, 1863 (24 spp.), Mey- tive to diameter of basal antennomeres (male) > 50%; rick, 1894 (8 spp.), Cybalobotys Maes, 2001 (3 spp.), 21:1, lobules on lateral edge of tympanal case present; Deltobotys Munroe, 1964 (3 spp.), Demobotys Munroe 54:2, depth of gap or split of juxta > 60% of dorsoventral & Mutuura, 1969 (2 spp.), Hübner, 1825 length of juxta to complete division into two juxta arms. (12 spp.), Hübner, 1818 (9 spp.), Epiparbattia Furthermore, Uresiphita shares with Portentomorphini Caradja, 1925 (2 spp.), Eumorphobotys Munroe & Mu- the elongate ediacaroid signum. Species of Uresiphita tuura, 1969 (2 spp.), Munroe, 1976 (1 sp.), mainly feed on Fabaceae (Munroe 1976a). Gynenomis Munroe & Mutuura, 1968 (2 spp.), Hahn­ Herpetobotys Maes, 2001 (3 spp.) shares the ediacar- cappsia Munroe, 1976 (39 spp.), Munroe, oid signum and the emergence of the appendix bursae at 1976 (5 spp.), Limbobotys Munroe & Mutuura, 1970 the anterior ductus bursae with Uresiphita, but the male (5 spp.), Munroeodes Amsel, 1957 (4 spp.), Nascia J. genitalia are different. For now, we leave Uresiphita and Curtis, 1835 (3 spp.), Neadeloides Klima, 1939 (2 spp.), Herpetobotys incertae sedis in Pyraustinae. Neoepicorsia Munroe, 1964 (7 spp.), Neohelvibotys The Australian species of Hyalobathra have been re- Munroe, 1976 (9 spp.), Nephelobotys Munroe & Mu- vised by Sutrisno & Horak (2003). tuura, 1970 (1 sp.), Motschulsky, 1861 (4 spp.), Oronomis Munroe & Mutuura, 1968 (1 sp.), Palepicor­ sia Maes, 1995 (1 sp.), Paranomis Munroe & Mutuura, 4.3.6. Pyraustini Meyrick, 1890 stat.rev. 1968 (4 spp.), Meyrick, 1890 (9 spp.), Par­ battia Moore, 1888 (6 spp.), Zeller, 1876 (1 Type genus: Pyrausta Schrank, 1802 = Botydes Blanchard, 1840 sp.), Placosaris Meyrick, 1897 (20 spp.), Powysia Maes, = Ennychites Duponchel, 1845 2006 (1 sp.), Prooedema Hampson, 1896 (1 sp.), Protepi­ corsia Munroe, 1964 (13 spp.), Pseudepicorsia Munroe, Synapomorphies. 9:0, third labial palpomere well de- 1964 (4 spp.), Pseudognathobotys Maes, 2001 (2 spp.), veloped in both sexes; 78:1, general shape of post-basal Pseudopagyda Slamka, 2013 (3 spp.), Pseudopolygram­ costa (not the entire dorsal valva edge) straight; 113:1, modes Munroe & Mutuura, 1969 (1 sp.), Pyrasia M. O. appendix bursae on corpus bursae present. Slow optimi- Martin, 1986 (1 sp.), Sarabotys Munroe, 1964 (2 spp.), zation only: 18:1, retinacular hook present in males. The Meyrick, 1890 (1 sp.), Sinibotys Munroe & parsimony trees imply a very different diagnosis: 5:0, Mutuura, 1969 (5 spp.), Thivolleo Maes, 2006 (4 spp.), basal antennomeres of male with sensillae ≤ 50% their Thliptoceras Warren, 1890 (31 spp.), Toxobotys Munroe diameter; 10:0, maxillary palpi long enough to contact & Mutuura, 1968 (3 spp.), Vittabotys Munroe & Mutuu- each other; 44:0, uncus conical, not capitate. ra, 1970 (1 sp.) and Xanthostege Munroe, 1976 (2 spp.). Description. The uncus is broad to elongate conical, Food plants. Achyra, Anania, Hyalorista, Loxostege without a prominent neck constriction and bulbous head; and Sitochroa are polyphagous on a variety of host the uncus has fne setae or robust bifurcate chaetae; many plants (Munroe 1976a; Robinson et al. 2010; Janzen taxa with a lobate process (sella sensu Marion 1952) on & Hallwachs 2009). The Central American species of the central inner valva, carrying long monoflament or Pyrausta mainly feed on Lamiaceae, Verbenaceae, Ama- multifd chaetae (editum sensu Marion 1952). The sig- ranthaceae (Janzen & Hallwachs 2009). Oenobotys is re- num is broad and rhombical, with the transverse axis corded from Eupatorium (Asteraceae) (Munroe 1976a).

193 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Pagyda species feed on Verbenaceae and Scrophulari- to Wurthiini based on our phylogenetic results. As in the aceae, Psammotis on Lamiaceae, and Pseudopyrausta studies of Mutanen et al. (2010) and Regier et al. (2012), on Lantana (Verbenaceae) (Robinson et al. 2010). The this taxon exhibits a very long terminal branch in our monotypic Paracorsia in mainly found on Fabaceae phylogenetic results (Fig. 1). The RogueNaRok analysis (Genista, Ulex, Cytisus, Phaseolus), but also on Scrophu- marked Niphopyralis as rogue taxon, but we decided to lariaceae (Verbascum) (Lhomme 1935). keep it in the dataset as we wanted to investigate its rela- Remarks. Our concept of Pyraustini still comprises tionship with other Spilomelinae (see 3.3.). Most of the the majority of the genera and species of Pyraustinae. observed substitutions in N. chionesis relative to other Although we do not further subdivide the tribe, many investigated taxa are synonymous, i.e. they do not cause characters would provide good evidence. For example, a change in the translated amino acid. the spatulate scales of the editum characterize many gen- We fnd little congruence between our phylogenetic era. Achyra, Loxostege, Powysia and Sitochroa share a results and those of the study by Sutrisno (2002a) based unique apomorphic anterior directed projection medially on 42 external and genital characters of adult moths of on the frons (1:1). Species of Anania share a unique syn- selected Australian Spilomelinae, partly due to the little apomorphic cone-shaped central structure in the antrum taxon overlap between the two datasets. Sutrisno (2002a) (97:1) (Leraut 2005a; Tränkner et al. 2009). proposed two synapomorphies for Spilomelinae, namely a strongly bilobed praecinctorium and the absence of a retinacular hook. We fnd the former character to not be consistent among Spilomelinae; the latter character is in- 5. Discussion deed not found among Spilomelinae, but present in most Pyraustinae, although it is reduced in many taxa. The common fndings in both phylogenies are: Isocentris + The phylogenetic analysis of our dataset results in the Hyalobathra, which we place in Portentomorphini; Hy­ same relationships among Crambidae as found by Regier menia + Spoladea, both in Hymeniini; Hymenoptychis + et al. (2012) based on a different set of molecular markers Tatobotys (misspelled as ‘Tatabotys’ in Sutrisno 2002a) that overlaps with our dataset in part of the CAD gene. are placed in Steniini. Furthermore, Agrioglypta, Chryso­ Because of the low number of Crambidae outgroup taxa thyridia, Didymostoma, Dysallacta, Glyphodes, in our dataset, comparison with the topology of Regier et and Talanga form a monophylum in Sutrisno (2002a), al. (2012) is only possible to a limited degree, but our re- and we place all seven genera in Margaroniini. We also sults refect their fndings where the “Wet Habitat Clade” fnd the synapomorphies proposed by Sutrisno (2002a) is sister to the clade of Crambinae + Scopariinae. among our synapomorphies for this clade, i.e. a heterog- regier et al. (2012) did not include a representative enous sclerotization of male tergite 8 (character 33:1), of the Sufetula genus group in their analysis. We include and two signa (character 108:1). Sufetula in our dataset, and we fnd it to fall outside of Our phylogenetic results largely refect those of the Spilomelinae, agreeing Hayden (2013) and Minet (2015) study of Haines & Rubinoff (2012) on Omiodes. In their who argued for the exclusion of the Sufetula group from phylogram (fg. 2 therein), nine of our proposed Spilo- Spilomelinae. Instead, Sufetula is sister to the “CAMMSS melinae tribes can be identifed: Udeini (Udea), Agroter- clade” minus sensu Regier et al. (2012). ini (Patania, Pleuroptya), Spilomelini (Cnaphalocrocis, Minet (2015) re-established the name Lathrotelinae on species misplaced in ‘Phostria’), Hymeniini (Spoladea), a subfamily rank for the Sufetula group and placed the Herpetogrammatini (Herpetogramma), Trichaeini (Pro­ taxon near Acentropinae based on shared characters of phantis), Nomophilini (Nomophila, Sameodes), Steniini the sternum on abdominal segment A2 and the oviposi- (Bradina, Piletocera), and Margaroniini (‘Omiodes’ ba­ tor. A phylogenetic analysis of a larger taxon sampling of salticalis and its sister clade). Interestingly, Haines & Crambidae, including all currently accepted subfamilies, Rubinoff (2012) found a clade comprising the still un- is necessary to investigate the relationship of Lathroteli- placed genera Prorodes, Syllepte and Coptobasis as sister nae within Crambidae. In his morphology-based phylo- to Bradina + Piletocera. genetic analysis of Australian Spilomelinae, Sutrisno The GENES- and TIGER-partitioned results differ (2002a) fnds Diplopseustis, now in Lathrotelinae (Minet in topology, the most fundamental difference being the 2015), as subordinate in Spilomelinae, and sister to the placement of Spilomelini, which in the GENES-parti- monotypic Aboetheta. tioned analyses is sister to Margaroniini (with PP << 0.9; Wurthiinae, with the single genus Niphopyralis (= see dotted line in Fig. 1). Other differences include the Wurthia), was originally described in Arctiidae (Roepke lack of support (i.e. PP < 0.9) in TIGER-partitioned anal- 1916). Kemner (1923) synonymised the group with Sch- yses for the monophyla Midila + Schoenobius, (Psammo­ oenobiinae, where Munroe (1958) retained it, while tis + Pseudopyrausta + Anania), and Udeini + Lineodini. Lewvanich (1981) transferred it to Pyraustinae (s.l.). Re- wahlberg et al. (2005) reported synergistic effects gier et al. (2012) found Niphopyralis to be ingroup of of combined morphological and molecular data for their a strongly supported Spilomelinae, and consequently phylogenetic analysis of Papilionoidea, and in their re- synonymized Wurthiinae with Spilomelinae. We confrm view of studies using these two kinds of data, Wortley that Niphopyralis belongs in Spilomelinae and assign it & Scotland (2006) fnd that most often node resolution

194 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

and support increase with the addition of morphological natural lines”, i.e. Spilomelinae and Pyraustinae sensu data to a genetic dataset. In our results, the phylogram is stricto. We concur with Forbes (1926), and furthermore nearly fully resolved, and most nodes have high support consider the presence of a retinacular hook the plesio- even in the dataset only comprising the molecular data morphic character state as it is found in a number of taxa (see posterior probabilities below branches in Fig. 1). in the sister group of Spilomelinae + Pyraustinae, e.g. in The only signifcant exception is within the Margaroniini Scopariinae (Nuss 2005), Crambinae (Landry 1995) and clade, where resolution is poor, and observed relation- Schoenobiinae (Lewvanich 1981); see Sauter (1973) for ships as well as their support vary across the differently a detailed study on this character among Pyraloidea. The partitioned analyses. In contrast to previous authors, the retinacular hook is absent in all investigated Spilomeli- addition of morphological data to our molecular dataset nae, and it may therefore serve as a diagnostic character does not result in an increase in topology resolution or for the group. In other subfamilies, however, the pres- branch support. ence or absence of this structure in males is highly vari- Based on the poor performance of preliminary anal- able at the generic (Solis & Maes 2003) or species level yses which included the morphological coding of the (Nuss 2005). Crambidae outgroup, we decided to omit this part of the Character 115 (locality of larval feeding) is the only data. The problems mainly concerned the convergence character of the immature life stage, and the only char- of the parallel MrBayes runs and resulted in ESS < 100 acter not concerning morphology. We chose to include for several parameters and a somewhat different topol- this character as we considered it as potentially carry- ogy, where Pyraustinae is sister to a monophylum of ing phylogenetic information. Although data coverage Spilomelinae and the Crambidae outgroup. Due to the for this character is only about 40%, some statements insuffcient ESS, this alternative topology was rejected. can be made from the data: For most taxa, there is no Furthermore, the choice and circumscription of the mor- apparent association between phylogenetic lineage and phological characters focus on Spilomelinae and Pyraus- larval feeding locality. The majority of coded taxa has tinae, and for many characters, we are not confdent about larvae that feed concealed in rolled or spun leaves or in drawing homologies with other groups of Crambidae. At a web (character state 115:0). This is the main feeding the same time, other crambids exhibit characters that locality for Margaroniini larvae, although some (Agath­ are not present in Spilomelinae and Pyraustinae, e.g. the odes, Liopasia, Maruca, Terastia) are partly or entirely well-developed gnathos of Scopariinae, Crambinae, Sch- borers in stems, branches, fowers, pods and/or fruits. An oenobiinae, Glaphyriinae and other groups. It is unlikely interesting association of potential phylogenetic value is to get meaningful results for the phylogeny of Spilomeli- the feeding of Dolicharthria larvae on decaying or dead nae if morphology is coded, for example, based on the plant matter (115:5), a behaviour that is observed in other characters defned by Landry (1995) for Crambini, or by Steniini as well (see ‘Food plants’ in 4.2.18.). Sutrisno (2002b) for the Australian Glyphodes species Larval host plants and feeding modes (internal vs. and resembling genera. Consequently, a morphomatrix external; leaf rolling, leaf webbing etc.) are considered that covers characters from all Crambidae taxa and that a useful source for future research on phylogenetic rela- is based on a less biased taxon sampling would be neces- tionships among Spilomelinae and Pyraloidea in general. sary to better refect the morphological diversity of the A study by Segar et al. (2017) on phylogenetic predic- focus group and to lead to more meaningful phylogenetic tions of host plant use in Pyraloidea and Geometridae results. found that host plant preference is phylogenetically rela- In contrast to the other excluded Crambidae outgroup tively conserved in snout moths. The host plants associa- taxa, we choose to retain the Lathrotelinae Sufetula in the tions of the taxa studied here support this observation, fnal morphological data matrix in order to investigate its e.g. Lineodini almost exclusively feeding on Solanaceae, placement in the phylogeny based on all available data. Asciodini commonly on Caryophyllales, and most Por- Sufetula was recently removed from Spilomelinae (Mi- tentomorphini on Phyllanthaceae. net 2015), and we concur with this decision as we fnd The majority of investigated Pyraustinae genera is the genus to not belong to Spilomelinae in our phyloge- placed in Pyraustini, a rather homogenous group which netic results. mainly varies in the shape of the valva and of the sclero- Pyraustinae and Spilomelinae are both strongly sup- tised processes on the inner valva surface, as well as in ported monophyletic and sister to each other, as found the uncus shape, although the spectrum of uncus varia- by Regier et al. (2012), but opposed to Solis & Maes tion is far narrower than in Spilomelinae. Munroe (1995) (2003) who found the two groups distantly related. The was uncertain about the inclusion of the Neotropical difference in the structure of the fornix tympani, recessed Portentomorpha and the related Old World Hyalobathra within the tympanic frame in Pyraustinae and projecting into his concept of Pyraustinae (Pyraustini sensu Munroe ventrally beyond the tympanic frame in Spilomelinae, is 1995). In our phylogenetic results, both genera are part the most consistent character for distinguishing the two of Pyraustinae, and they are placed in Portentomorphini. groups, and underlines the importance of the tympana However, we fnd Hyalobathra to be closer related to the for pyraloid systematics. Forbes (1926: p. 332) mentions African Cryptosara. Tetridia, the sister to all other phylo- that the absence or presence of the retinacular hook sep- genetically investigated Pyraustinae, requires taxonomi- arates Pyraustinae (s.l.) into two “mainly if not wholly cal revision (see Systematics under 4.3.1.).

195 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Spilomelinae is found highly supported monophyl- The poorly resolved relationships among Margaronii- etic in our analysis, supporting Munroe’s (1995) opin- ni should be addressed through the choice of better-suited ion of Spilomelinae being “at least in large part” mono- genetic markers, e.g. DDC which has a substitution rate phyletic, while contradicting Minet (1982) and Solis & similar to COI (Wahlberg & Wheat 2008). Furthermore, Maes (2003) who perceived Spilomelinae as a para- or the morphological dataset can be improved to incorpo- polyphyletic assemblage. The only uniquely derived apo- rate additional characters, e.g. structure of the hairpencil morphy common to all investigated Spilomelinae is the pads and scales, shape and sclerotisation patterns of the ventrad projecting fornix tympani, a character present in valvae, and shape and structure of the tegumen and un- Minet’s (1982) circumscription of the group. A (strong- cus. ly) bilobed praecinctorium, considered diagnostic by Mi- With the morphological dataset at hand and the phy- net (1982) and Sutrisno (2002a), is found to be homo- logenetic results, we start to gain a better understanding plastic. It is absent in several investigated Spilomelinae of functional morphology of the genitalia and character and present in a number of Pyraustinae, and therefore of correlations. In most investigated Spilomelinae, the pres- no diagnostic use. According to Solis & Maes (2003) a ence of sclerotised membraneous strips on the pleural bilobed praecinctorium is also present in Midilinae. Ab- membranes of the male abdominal segment 8 corresponds sence characters like the lack of chaetosemata, the sub- with the presence of hairpencils, and we assume that the costal retinacular hook, and a well-developed gnathos in strips might serve as muscular attachment sites for retrac- males, as well as the large rhombical signum in females tion of the hairpencils. In Pyraustinae, this correlation is are not exclusive for Spilomelinae. The loss of the reti- absent, and we fnd sclerites on the pleurites of the male nacular hook is a synapomorphy of Spilomelinae, but it is segment 8 only in Euclasta, although most pyraustines paralleled in some Pyraustinae. The character of distinct- exhibit ‘simple’ hairpencils, i.e. a single small hairpencil ly tapered spinulae (Minet 1982) was not investigated. pad on each side of the anterior vinculum-tegumen con- According to Allyson (1981, 1984), there are no diag- nection, studded with one type of hairpencil scales. The nostic morphological characters distinguishing larvae of longitudinal membranous strip in the female genitalia’s Pyraustinae from those of Spilomelinae. antrum possibly functions as a stretching zone during In our phylogenetic results, the tribe Margaroniini copulation, when the male transfers the spermatophore. comprises the most sampled species. This could be due This character is present in species with narrow antrum to a sampling bias, but the large number of taxa attributed while it is rarely found in species with a broad antrum; to the tribe (currently 67 genera with 1,044 species) sup- it occurs in both Spilomelinae and Pyraustinae. We fnd ports this view of Margaroniini being the most diverse the fbula and the distal sacculus often in close spatial as- clade in Spilomelinae. Margaroniini is predominantly sociation, and the ventral valva margin is often less scle- tropical and subtropical in distribution. Apart from com- rotised in this area, suggesting that this complex might prising a large number of genera, the tribe also contains function as a bendable joint during copulation, when the many species-rich genera, like Palpita, Glyphodes, Omi­ male clasps the female genital with its valvae. Minet et odes and Diaphania. al. (2014) identifed the same weak fexure on the saccu- We fnd several well-supported relationships among lus as a character of Noctuidae s. str. including Dyopsi- Margaroniini: the larvae of Liopasia, Terastia and Ag­ nae. In the future, emphasis should be put on studying athodes all feed on Erythrina, and imagines of the three the muscular attachment regions of this supposed point moth genera share a similar wing pattern (Sourakov et for valval bending, as well as of the vinculum and tegu- al. 2015). We fnd the three genera to be closely related, men (see Kuznetzov & Stekolnikov 1979a,b). Like for with Liopasia being sister to Agathodes + Terastia, as re- most other characters studied, investigation of muscula- ported by Sourakov et al. (2015). The Glyphodes group, ture and nervature is essential to better understand their as circumscribed by Sutrisno (2002b) and Sutrisno et function and homology. Furthermore, the ontogenetic al. (2006), further includes Obtusipalpis as well as the origins of the fbula should be investigated as it is not Dichocrocis zebralis species complex. Conogethes pan­ clear whether the sclerotised protrusions on the inner side damalis, related to the species complex of the yellow of the valva that are found in most species of Spilomeli- peach moth C. punctiferalis, is found to be sister to the nae are homologous. We assume that protrusions emerg- Neotropical genus Azochis. They share a similar anatomy ing from the costa are not homologous with those arising of the uncus, tegumen and valvae, and a similar structure from the central inner valva, and we consequently code of the hairpencil scales. The phylogenetic relationships them as separate characters. to species with a highly similar wing pattern, like the The morphological circumscription of the observed African Marwitzia species (Maes 1998b) or species of tribes in Spilomelinae and Pyraustinae allows for the as- the Neotropical Polygrammodes eleuata complex, is not signment of additional taxa in those tribes through mor- known. phological investigation, without the strict requirement The sister group of Omiodes, as investigated by of molecular data. These morphological diagnoses allow Haines & Rubinoff (2012), is still not known due to the the assignment of additional 125 genera to Spilomelinae extensive polytomy in Margaroniini. We can, however, tribes, and additional 56 genera to Pyraustinae tribes. 135 rule out Cnaphalocrocis as sister group, which we place genera of Spilomelinae and 103 genera of Pyraustinae re- in Spilomelini. main unassigned to any of the proposed tribes. Among the

196 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

Table 3. Correlation of Munroe’s (1995) proposed Spilomelinae genus groups to the Spilomelinae tribes proposed herein.

Munroe‘s (1995) genus groups proposed tribes unplaced genera other family groups Phaedropsis group Agroterini Syllepte group Agroterini Syngropia, Praephostria, Syllepte, Troctoceras Herpetogramma group Herpetogrammatini Pelinopsis Hydriris group Hydririni apart from Geshna (Spilomelini) Udea group Lineodini apart from Lamprosema (Hydri­rini) and Udea (Udeini) Hymenia group Ercta: Udeini; Spilomela: Spilomelini; Blepharo­ Sacculosia mastix: Herpetogrammatini; Hymenia, Spoladea: Hymeniini; Anageshna, Apogeshna, Duponchelia, Loxostegopsis, Parastenia (= Dolicharthria), Penestola, Steniodes: Steniini; Desmia, Diasemiodes, Diasemiop­ sis, Diathrausta: Nomophilini Diaphania group Margaroniini Chromodes Polygrammodes group Margaroniini Siga group Cirrhocephalina, Siga, Zeuzerobotys: Spilomelini; Beebea, Laniifera, Laniipriva: Asciodini Eulepte group Gonocausta, Ommatospila, Syllepis: Hydri­rini Eulepte, Praeacrospila, Leucochromodes, Mesocondyla Samea group Nomophilini Stenorista Psara group Asciodini Conchylodes group Conchylodes: Udeini Pycnarmon Syngamia group Marasmia, Salbia: Spilomelini; Sufetula to Lathrotelinae (Minet Syngamia: Nomophilini 2015) Diaphantania group Wurthiini unplaced genera Nonazochis (= Conchylodes), Sisyracera, Tanaophysa: Agrammia, Carthade, Coelorhyncidia, Co­ Ennomosia to Glaphyriinae (Hayden Udeini; Bocchoropsis, Coenostolopsis, Cyclocena remata, , Deuterophysa, Dichocrocis, & Dickel 2014); Hydropionea and (Hayden & Dickel 2014: syn. of Microthyris), Gypo­des: , Heterudea, Hyalea, Ischnurg­ Plantegumia to Glaphyriinae; Agroterini; Analyta, Caprinia, Meroctena, Tyspa­nodes: es, , , , , Orthoraphis to Lathrotelinae Margaroniini; Palpusia, Rhectocraspeda: Spilomelini; Metraeopsis, Microphysetica, Mimudea, Eurrhyparodes: Herpetogrammatini; Bicilia, Loxomor­ Pectinobotys, Piletocera, Piletosoma, Platyg­ pha, Maracayia: Asciodini; Trichaea: Trichaeini; Bra­ raphis, Plectrona, Syntrita, Tanao­physopsis, dina: Steniini; Bocchoris, Parapilocrocis: Nomophilini Trithyris unassigned Spilomelinae genera, the most species-rich taxa that do not ft into this system of tribes. Such taxa are Syllepte (198 spp.), Nacoleia (84 spp.), Pycnarmon can be morphologically coded as well as sequenced for (59 spp.), Dichocrocis (53 spp.) and Mimudea Warren, the six genetic markers used in this study. Their phyloge- 1892 (42 spp.), while 76 genera contain only one to two netic placement can then be inferred through a combina- species, respectively. In Pyraustinae, the most species- tion with the data presented here. We therefore provide a rich unassigned genera are Semniomima Warren, 1892 ‘modular’ dataset where taxa of interest can be added in (15 spp.), Calamochrous Lederer, 1863 (13 spp.) and order to refne the circumscription of the proposed tribes Moore, 1886 (12 spp.). In the European fauna of and to widen our understanding of the phylogenetic rela- Spilomelinae and Pyraustinae, only Uresiphita remains tionships of Spilomelinae and Pyraustinae. The resulting unplaced as sister to Portentomorphini. In the Nearctic improved understanding of Spilomelinae and Pyrausti- region, Daulia, Deuterophysa, Eulepte, Microphysetica, nae genera is expected to promote taxonomic revisions Stenochora and Syllepte are still unplaced. Furthermore, of genera and species groups as well as ecological and the majority of genera from Munroe’s (1995) Neotropi- applied research on the pyraloids. cal genus groups are placed in tribes (Table 3). Ultimately, the type species of every spilomeline and pyraustine genus should be investigated and placed into the phylogenetic framework. The morphological 6. Acknowledgements circumscription of the observed tribes in Spilomelinae and Pyraustinae allows for the assignment of additional We are grateful to the following colleagues for providing material taxa in those tribes through morphological investigation, and data for our study: Leif Aarvik (Natural History Museum Oslo, without the strict requirement of molecular data. This Norway), Stoyan Beshkov (National Museum of Natural History should be applied to the 132 Spilomelinae genera and Sofa, Bulgaria), Jason Dombroskie (Cornell University Ithaca, New 103 Pyraustinae genera which are not yet assigned to any York, U.S.A.), Markus Franzén (Helmholtz Centre for Environmen- of the tribes. A concerted effort among systematists to tal Research Leipzig, Germany), Will Haines (University of Hawaii at Manoa, Hawaii, USA), John B. Heppner, Jacqueline Y. Miller morphologically investigate those unplaced genera and and Deborah Matthews (all McGuire Center for Lepidoptera & to assign them to the proposed tribes is feasible and de- , Gainesville, Florida, USA), Marianne Horak (CSIRO sirable. This effort will likely result in the recognition of Australia), Timm Karisch (Museum of Natural History Dessau,

197 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

Germany), Bernard Landry (Natural History Museum Geneva, Chen K., Horak M., Du X., Zhang D.-D. 2017. Revision of the Switzerland), Knud Larsen (Dyssegård, ), Norbert Pöll Australian species of Agrotera Schrank (Lepidoptera: Pyraloi­ (Biologiezentrum Linz, Austria), John Rawlins (Carnegie Museum dea: Crambidae: Spilomelinae). – Zootaxa 4362(2): 213 – 224. of Natural History, Pittsburgh, USA), Rodolphe Rougerie (Natural Clarke J.F.G. 1986. Pyralidae and of the Mar- History Museum Paris, France), Andreas Segerer (Zoological State quesas Archipelago. – Smithsonian Contributions to Zoology Collections Munich, Germany), Stephen Sutton and Charles Vairap- 416: 1 – 485. pan (University of Sabah, Kota Kinabalu, Malaysia), Marja van der Clavijo Albertos J.A. 1990. Systematics of black and white spe- Straten (Nederlandse Voedsel- en Warenautoriteit Wageningen, The cies of the genus Diaphania Hübner, 1818 (Lepidoptera: Pyrali- ), Boyan Zlatkov (Sofa University “St. Kliment Ohrid- dae: Pyraustinae). – Dissertation, Department of Entomology, ski” Sofa, Bulgaria) and Andreas Zwick (Australian National McGill University, Montreal, Canada. xi + 215 pp., 159 fgs. Collection, CSIRO, Canberra, Australia), and especially the late Common I.F.B. 1990. Moths of Australia. – Melbourne University Linwood C. Dow (Largo, Florida), whose efforts to collect, curate, Press: Carlton, Victoria, vi + 535 pp. and professionally identify Neotropical pyraloids was especially Cummins C.A., McInerney J.O. 2011. A method for inferring the useful for making genera available to study through his bequest. rate of evolution of homologous characters that can potentially We are thankful to Miss Chung (Sabah Biodiversity Centre, improve phylogenetic inference, resolve deep divergence and Malaysia), Prof. Charles S. Vairappan and Stephen Sutton (both correct systematic biases. – Systematic Biology 60(6): 833 – 844. University Malaysia Sabah, Malaysia), and the staff of Sabah Parks Darriba D., Taboada G.L., Doallo R., Posada D. 2012. jModel- and the University Malaysia Sabah for collecting preparations Test 2: more models, new heuristics and parallel computing. – and support in Mount Kinabalu National Park. We further thank Nature Methods 9(8): 772. Manuela Bartel, Anke Müller and Anja Rauh (all Senckenberg de Prins W., Mazzei P. 2016. Some faunistic notes on selected moth Natural History Collections Dresden, Germany) as well as Louise species (Lepidoptera) from the . – Phelsuma 24: 21 – 34. 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DAMBE5: a comprehensive software package for Guenée and allied genera (Lepidoptera: Crambidae; Spilomeli- data analysis in molecular biology and evolution. – Molecular nae). – Entomological Science 5(4): 457 – 467. Biology and Evolution 30(7): 1720 – 1728. Sutrisno H. 2003. Phylogeny of Glyphodes Guenée (Lepidoptera: Xia X., Lemey P. 2009. Assessing substitution saturation with Crambidae: Spilomelinae) based on nucleotide sequence varia- DAMBE. Pp. 615 – 630 in: Lemey P., Salemi M., Vandamme A.- tion in mitochondrial CO I gene: congruence with morphological M. (eds), The Phylogenetic Handbook: A Practical Approach to data. – Treubia 33(1): 35 – 42. DNA and Protein Phylogeny. 2nd edition. – Cambridge Univer- Sutrisno H. 2004. Phylogeny of the two closely-related moth ge­ sity Press. nera, Agrioglypta Meyrick and Talanga Moore (Lepidoptera: Xia X., Xie Z., Salemi M., Chen L., Wang Y. 2003. An index of Cram­bidae; Spilomelinae) based on morphology and mitochon- substitution saturation and its application. – Molecular Phyloge- drial COII sequence variations. – Hayati. Journal of Biosciences netics and Evolution 26: 1 – 7. 11: 93 – 97. Zimmerman E.C. 1958. Lepidoptera: Pyraloidea. – Insects of Ha- Sutrisno H. 2005. Molecular phylogeny of Agrioglypta Meyrick waii 8: i – xii, 1 – 456. and Talanga Moore (Lepidoptera: Crambidae; Spilomelinae) in- ferred from nuclear EF-1a gene. – Hayati. Journal of Biosciences 12(2): 45 – 49. Sutrisno H. 2006. Evolution of a wingless gene and its utility for 8. Appendix inferring the relationships within Glyphodes moths. – Hayati. Journal of Biosciences 13(4): 145 – 150. Sutrisno H., Azuma N., Higashi S. 2006. Molecular phylogeny of Checklist of Spilomelinae and Pyraustinae tribes, associ- the Indo-Australian Glyphodes and its allied genera (Insecta: ated genera and species numbers: Lepidoptera: Crambidae: Spilomelinae) inferred from mitochon- drial COI and COII and nuclear EF-1α gene sequences. – Spe- SPILOMELINAE cies Diversity 11: 57 – 69. Sutrisno H., Horak M. 2003. Revision of the Australian species Hydririni: Choristostigma Warren, 1892 (10 spp.) — Gonocaus­ of Hyalobathra Meyrick (Lepidoptera: Pyraloidea: Crambidae: ta Lederer, 1863 (4 spp.) — Hydriris Meyrick, 1885 (7 spp.) — Pyraustinae) based on adult morphology and with description Lamprosema Hübner, 1823 (72 spp.) [polyphyletic] — Nehydriris of a new species. – Australian Journal of Entomology 42(3): Munroe, 1974 (1 sp.) — Ommatospila Lederer, 1963 (3 spp.) — 233 – 248. Rhectothyris Warren, 1890 (1 sp.) — Syllepis Poey, 1832 (7 spp.) Swofford D.L., Maddison W.P. 1987. Reconstructing ancestral Lineodini: Atomopteryx Walsingham, 1891 (10 spp.) — Euleuci­ char­acter states under Wagner parsimony. – Mathematical Bio- nodes Capps, 1948 (1 sp.) — Leucinodes Guenée, 1854 (20 spp.) sciences 87: 199 – 229. [MALLY et al. 2015; misplaced spp. in Asia and Australia] — Li­ Tominaga S. 2002. Biological notes on fve species of the Pyra- neodes Guenée, 1854 (39 spp.) — Neoleucinodes Capps, 1948 loidea from Okinawa Island. – Yugato 167: 9 – 12. (9 spp.) — Proleucinodes Capps, 1948 (4 spp.) — Rhectosemia Tränkner A., Li H., Nuss M. 2009. On the systematics of Ana­ Lederer, 1863 (12 spp.) nia Hübner, 1823 (Pyraloidea: Crambidae: Pyraustinae). – Nota Udeini: Cheverella B. Landry, 2011 (1 sp.) — Conchylodes Lepidopterologica 32(1): 63 – 80. Guenée, 1854 (21 spp.) [paraphyletic?] — Deana Butler, 1879 (1 Ullah M., Yang Z., Qiao P., Zhang Y. 2017. A new cryptic species sp.) — Ercta Walker, 1859 (7 spp.) — Mnesictena Meyrick, 1884 of Nagiella Munroe from China revealed by DNA barcodes and (7 spp.) — Sisyracera Möschler, 1890 (3 spp.) — Tanaophysa War- morphological evidence. – ZooKeys 679: 65 – 76. ren, 1892 (2 spp.) — Udea Guenée, 1845 (in Duponchel) (214 spp.) Wagner M.R., Cobbinah J.R., Bosu P.P. 2008. Forest Entomology [e.g. MUNROE 1966, INOUE et al. 2008, MALLY & NUSS 2011] in West Tropical Africa: Forest Insects of . – Springer. lxx — Udeoides Maes, 2006 (5 spp.) + 224 pp. Wurthiini: Apilocrocis Amsel, 1956 (11 spp.) — Aristebulea Wahlberg N., Wheat C.W. 2008. Genomic outposts serve the Munroe & Mutuura, 1968 (2 spp.) — Diaphantania Möschler, phylogenomic pioneers: Designing novel nuclear markers for 1890 (3 spp.) — Mimetebulea Munroe & Mutuura, 1968 (1 sp.) genomic DNA extractions of Lepidoptera. – Systematic Biology — Niphopyralis Hampson, 1893 (8 spp.) — Pseudebulea Butler, London 57(2): 231 – 242. 1881 (4 spp.) Wahlberg N., Braby M.F., Brower A.V.Z., de Jong R., Lee M.-M., Agroterini: Aetholix Lederer, 1863 (4 spp.) — Agrotera Schrank, Nylin S., Pierce N.E., Sperling F.A.H., Vila R., Warren A.D., 1802 (24 spp.) [CHEN et al. 2017] — Aiyura Munroe, 1974 (2

202 ARTHROPOD SYSTEMATICS & PHYLOGENY — 77 (1) 2019

spp.) — Bocchoropsis Amsel, 1956 (2 spp.) — Chalcidoptera 1892 (2 spp.) — Salbia Guenée, 1854 (35 spp.) — Scaptesylodes Butler, 1887 (15 spp.) — Chilochromopsis Munroe, 1964 (1 sp.) Munroe, 1976 (2 spp.) — Siga Hübner, 1820 (2 spp.) — Spilomela — Coenostolopsis Munroe, 1960 (3 spp.) — Diastictis Hübner, Guenée, 1854 (8 spp.) — Zeuzerobotys Munroe, 1963 (1 sp.) 1818 (12 spp.) — Framinghamia Strand, 1920 (1 sp.) — Glau­ Herpetogrammatini: Blepharomastix Lederer, 1863 (85 spp.) cobotys Maes, 2008 (1 sp.) — Goliathodes Munroe, 1974 (1 sp.) — Cryptobotys Munroe, 1956 (2 spp.) — Eurrhyparodes Snel- — Gypodes Munroe, 1976 (1 sp.) — Haritalodes Warren, 1890 (11 len, 1880 (12 spp.) — Herpetogramma Lederer, 1863 (100 spp.) spp.) — Lygropia Lederer, 1863 (68 spp.) — Lypotigris Hübner, — Hileithia Snellen, 1875 (19 spp.) — Pilocrocis Lederer, 1863 1825 (1 sp.) — Micromartinia Amsel, 1957 (1 sp.) — Microthyris (65 spp.) Lederer, 1863 (7 spp.) — Nagiella Munroe, 1976 (4 spp.) — Neo­ Hymeniini: Hymenia Hübner, 1825 (3 spp.) — Spoladea Guenée, analthes Yamanaka & Kirpichnikova, 1993 (8 spp.) — Nosophora 1854 (2 spp.) Lederer, 1863 (26 spp.) — Notarcha Meyrick, 1884 (18 spp.) — Asciodini: Arthromastix Warren, 1890 (2 spp.) — Asciodes Pantographa Lederer, 1863 (9 spp.) — Patania Moore, 1888 (40 Guenée, 1854 (5 spp.) — Beebea Schaus, 1923 (1 sp.) — Bicilia spp.) — Phaedropsis Warren, 1890 (24 spp.) — Phostria Hübner, Amsel, 1956 (4 spp.) — Ceratocilia Amsel, 1956 (8 spp.) — Cera­ 1819 (87 spp.) — Phryganodes Guenée, 1854 (26 spp.) — Tetra­ toclasis Lederer, 1863 (9 spp.) — Erilusa Walker, 1866 (3 spp.) cona Meyrick, 1884 (2 spp.) [CHEN et al. 2017] — Ulopeza Zeller, [tentative placement] — Laniifera Hampson, 1899 (1 sp.) — Lani­ 1852 (16 spp.) ipriva Munroe, 1976 (1 sp.) — Loxomorpha Amsel, 1956 (4 spp.) Margaroniini: Agathodes Guenée, 1854 (16 spp.) — Agrioglypta — Maracayia Amsel, 1956 (2 spp.) — Psara Snellen, 1875 (36 Meyrick, 1932 (11 spp.) [SUTRISNO 2002a,b, 2005, SUTRISNO spp.) — Sathria Lederer, 1863 (3 spp.) et al. 2006] — Alytana J. C. Shaffer, & Munroe, 2007 (2 spp.) — Trichaeini: Prophantis Warren, 1896 (8 spp.) — Sacculosia Amsel, Analyta Lederer, 1863 (10 spp.) — Anarmodia Lederer, 1863 (24 1956 (1 sp.) — Trichaea Herrich-Schäffer, 1866 (11 spp.) — Ze­ spp.) — Antigastra Lederer, 1863 (2 spp.) — Aphytoceros Mey- namorpha Amsel, 1956 (2 spp.) rick, 1884 (3 spp.) — Arthroschista Hampson, 1893 (2 spp.) — Steniini: Anageshna Munroe, 1956 (1 sp.) — Apogeshna Munroe, Asturodes Amsel, 1956 (1 sp.) — Azochis Walker, 1859 (16 spp.) 1956 (3 spp.) — Bradina Lederer, 1863 (87 spp.) — Dolicharthria — Botyodes Guenée, 1854 (10 spp.) — Cadarena Moore, 1886 (1 Stephens, 1834 (24 spp.) — Duponchelia Zeller, 1847 (5 spp.) — sp.) — Caprinia Walker, 1859 (11 spp.) — Chabulina J. C. Shaffer, Epherema Snellen, 1892 (1 sp.) — Hymenoptychis Zeller, 1852 (4 & Munroe, 2007 (2 spp.) — Charitoprepes Warren, 1896 (2 spp.) spp.) — Loxostegopsis Dyar, 1917 (6 spp.) — Metasia Guenée, — Chrysophyllis Meyrick, 1934 (1 sp.) — Chrysothyridia Munroe, 1854 (88 spp.) — Penestola Möschler, 1890 (3 spp.) — Steniodes 1967 (2 spp.) — Cirrhochrista Lederer, 1863 (38 spp.) — Colo­ Snellen, 1875 (9 spp.) — Symmoracma Meyrick, 1894 (1 sp.) — mychus Munroe, 1956 (2 spp.) — Compacta Amsel, 1956 (4 spp.) Tatobotys Butler, 1881 (11 spp.) — Condylorrhiza Lederer, 1863 (4 spp.) — Conogethes Meyrick, Nomophilini: Arnia Guenée, 1849 (1 sp.) — Ategumia Amsel, 1884 (16 spp.) [e.g. INOUE & YAMANAKA 2006, SHASHANK 1956 (10 spp.) — Bocchoris Moore, 1885 (31 spp.) — Crocidoc­ et al. 2015, 2018] — Cydalima Lederer, 1863 (9 spp.) [STRELT- nemis Warren, 1889 (2 spp.) — Desmia Westwood, 1832 (89 spp.) ZOV 2008, MALLY & NUSS 2010] — Diaphania Hübner, 1818 — Diacme Warren, 1892 (10 spp.) — Diasemia Hübner, 1825 (95 spp.) [CLAVIJO ALBERTOS 1990] — Didymostoma Warren, (13 spp.) — Diasemiodes Munroe, 1957 (4 spp.) — Diasemiopsis 1892 (2 spp.) [SUTRISNO 2002a] spp.) — Dysallacta Lederer, Munroe, 1957 (2 spp.) — Diathrausta Lederer, 1863 (20 spp.) — 1863 (3 spp.) [SUTRISNO 2002a] — Endocrossis Meyrick, 1889 Epipagis Hübner, 1825 (14 spp.) — Mecyna Doubleday, 1849 (34 (4 spp.) — Eusabena Snellen, 1901 (4 spp.) — Filodes Guenée, spp.) — Mimophobetron Munroe, 1950 (1 sp.) — Mimorista War- 1854 (16 spp.) — Ghesquierellana Berger, 1955 (5 spp.) — ren, 1890 (15 spp.) — Niphograpta Warren, 1892 (1 sp.) — No­ Glyphodella J. C. Shaffer & Munroe, 2007 (3 spp.) — Glyphodes mophila Hübner, 1825 (14 spp.) [MUNROE 1973] — Nothomastix Guenée, 1854 (156 spp.) [SUTRISNO 2002a,b, 2003, 2006, Warren, 1890 (5 spp.) — Parapilocrocis Munroe, 1967 (3 spp.) SUTRISNO et al. 2006] — Hedyleptopsis Munroe, 1960 (1 sp.) — Pardomima Warren, 1890 (16 spp.) — Perisyntrocha Meyrick, — Heterocnephes Lederer, 1863 (4 spp.) — Hodebertia Leraut, 1894 (4 spp.) — Pessocosma Meyrick, 1884 (4 spp.) — Samea 2003 (1 sp.) — Hoterodes Guenée, 1854 (5 spp.) — Leucochroma Guenée, 1854 (28 spp.) — Sameodes Snellen, 1880 (15 spp.) — Guenée, 1854 (6 spp.) — Liopasia Möschler, 1882 (15 spp.) — Syngamia Guenée, 1854 (25 spp.) Loxmaionia Schaus, 1913 (1 sp.) — Maruca Walker, 1859 (4 spp.) — Marwitzia Gaede, 1917 (3 spp.) — Megaphysa Guenée, 1854 (1 sp.) — Megastes Guenée, 1854 (16 spp.) — Meroctena Lederer, PYRAUSTINAE 1863 (4 spp.) — Nolckenia Snellen, 1875 (1 spp.) — Obtusipalpis Hampson, 1896 (6 spp.) — Omiodes Guenée, 1854 (98 spp.) [poly- Euclastini: Euclasta Lederer, 1855 (17 spp.) [POPESCU-GORJ & phyletic, HAINES & RUBINOFF 2012] — Omphisa Moore, 1886 CONSTANTINESCU 1977] (10 spp.) — Pachynoa Lederer, 1863 (12 spp.) — Palpita Hübner, Portentomorphini: Cryptosara E. L. Martin in Marion, 1957 (3 1808 (162 spp.) [INOUE 1996, 1997, 1999] — Parotis Hübner, spp.) — Hyalobathra Meyrick, 1885 (21 spp.) [SUTRISNO & 1831 (37 spp.) — Poliobotys J. C. Shaffer & Munroe, 2007 (1 HORAK 2003] — Isocentris Meyrick, 1887 (7 spp.) — Pionea­ sp.) — Polygrammodes Guenée, 1854 (78 spp.) — Polygrammop­ bathra J. C Shaffer & Munroe, 2007 (1 sp.) — Portentomorpha sis Munroe, 1960 (1 sp.) — Prenesta Snellen, 1875 (18 spp.) — Amsel, 1956 (1 sp.) Pygospila Guenée, 1854 (10 spp.) — Radessa Munroe, 1977 (2 Pyraustini: Achyra Guenée, 1849 (19 spp.) — Adoxobotys Mun- spp.) — Rhagoba Moore, 1888 (2 spp.) — Rhimphalea Lederer, roe, 1978 (3 spp.) — Aglaops Warren, 1892 (4 spp.) — Anamalaia 1863 (12 spp.) — Sinomphisa Munroe, 1958 (3 spp.) — Sparagmia Munroe & Mutuura, 1969 (1 sp.) — Anania Hübner, 1823 (117 Guenée, 1854 (1 sp.) — Stemorrhages Lederer, 1863 (8 spp.) — spp.) — Arenochroa Munroe, 1976 (1 sp.) — Aurorobotys Mun- Synclera Lederer, 1863 (13 spp.) — Syngamilyta Strand, 1920 (5 roe & Mutuura, 1971 (2 spp.) — Callibotys Munroe & Mutuura, spp.) — Talanga Moore, 1885 (9 spp.) [SUTRISNO 2002a,b, 2005, 1969 (3 spp.) — Carminibotys Munroe & Mutuura, 1971 (1 sp.) SUTRISNO et al. 2006] — Terastia Guenée, 1854 (7 spp.) — Tes­ — Ceuthobotys Munroe, 1978 (1 sp.) — Chilochroma Amsel, 1956 sema J. F. G. Clarke, 1986 (1 sp.) — Tyspanodes Warren, 1891 (4 spp.) — Chilocorsia Munroe, 1964 (1 sp.) — Chilopionea Mun- (20 spp.) — Uncobotyodes Kirti & Rose, 1990 (1 sp.) — Zebronia roe, 1964 (1 sp.) — Circobotys Butler, 1879 (19 spp.) — Croci­ Hübner, 1821 (6 spp.) dophora Lederer, 1863 (24 spp.) — Crypsiptya Meyrick, 1894 (8 Spilomelini: Aethaloessa Lederer, 1863 (3 spp.) — Cirrhocephali­ spp.) — Cybalobotys Maes, 2001(3 spp.) — Deltobotys Munroe, na Munroe, 1995 (5 spp.) — Cnaphalocrocis Lederer, 1863 (27 1964 (3 spp.) — Demobotys Munroe & Mutuura, 1969 (2 spp.) spp.) — Eporidia Walker, 1859 (1 sp.) — Geshna Dyar, 1906 (1 — Ecpyrrhorrhoe Hübner, 1825 (12 spp.) — Epicorsia Hübner, sp.) — Marasmia Lederer, 1863 (9 spp.) — Marasmianympha 1818 (9 spp.) — Epiparbattia Caradja, 1925 (2 spp.) — Eumor­ Munroe, 1991 (1 sp.) — Orphanostigma Warren, 1890 (6 spp.) phobotys Munroe & Mutuura, 1969 (2 spp.) — Fumibotys Mun- — Palpusia Amsel, 1956 (10 spp.) — Rhectocraspeda Warren, roe, 1976 (1 sp.) — Gynenomis Munroe & Mutuura, 1968 (2 spp.)

203 Mally et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

— Munroe, 1976 (39 spp.) — Helvibotys Munroe, Maes, 2006 (1 sp.) — Prooedema Hampson, 1891 (1 sp.) — Pro­ 1976 (5 spp.) — Hyalorista Warren, 1892 (5 spp.) — Limbobo­ tepicorsia Munroe, 1964 (13 spp.) — Psammotis Hübner, 1825 (8 tys Munroe & Mutuura, 1970 (5 spp.) — Loxostege Hübner, 1825 spp.) — Pseudepicorsia Munroe, 1964 (4 spp.) — Pseudognatho­ (90 spp.) — Munroeodes Amsel, 1957 (4 spp.) — Nascia J. Cur- botys Maes, 2001 (2 spp.) — Pseudopagyda Slamka, 2013 (3 spp.) tis, 1835 (3 spp.) — Neadeloides Klima, 1939 (2 spp.) — Neoepi­ [CHEN & ZHANG 2017] — Pseudopolygrammodes Munroe & corsia Munroe, 1964 (7 spp.) — Neohelvibotys Munroe, 1976 (9 Mutuura, 1969 (1 sp.) — Pseudopyrausta Amsel, 1956 (6 spp.) — spp.) — Nephelobotys Munroe & Mutuura, 1970 (1 sp.) — Nomis Pyrasia M. O. Martin, 1986 (1 sp.) — Pyrausta Schrank, 1802 (341 Motschulsky, 1861 (4 spp.) — Oenobotys Munroe, 1976 (5 spp.) spp.) — Sarabotys Munroe, 1964 (2 spp.) — Sclerocona Meyrick, — Oronomis Munroe & Mutuura, 1968 (1 sp.) — Ostrinia Hübner, 1890 (1 sp.) — Sinibotys Munroe & Mutuura, 1969 (5 spp.) — 1825 (21 spp.) — Pagyda Walker, 1859 (26 spp.) — Palepicorsia Sitochroa Hübner, 1825 (10 spp.) — Thivolleo Maes, 2006 (4 spp.) Maes, 1995 (1 sp.) — Paracorsia Marion, 1959 (1 sp.) — Para­ — Thliptoceras Warren, 1890 (31 spp.) — Toxobotys Munroe & nomis Munroe & Mutuura, 1968 (4 spp.) — Paratalanta Meyrick, Mutuura, 1968 (3 spp.) — Vittabotys Munroe & Mutuura, 1970 1890 (9 spp.) — Parbattia Moore, 1888 (6 spp.) — Perispasta Zel- (1 sp.) — Xanthostege Munroe, 1976 (2 spp.) ler, 1876 (1 sp.) — Placosaris Meyrick, 1897 (20 spp.) — Powysia

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File 1: mally&al-spilomelinaephylogeny-asp2019-electronic Present article: http://zoobank.org/urn:lsid:zoobank. supplement-1.xlsx — Table S1. List of examined genitalia slides. org:pub:07A92FE2-FB09-4216-A198-0CE041B86702 — DOI: 10.26049/ASP77-1-2019-07/1

File 2: mally&al-spilomelinaephylogeny-asp2019-electronic supplement-2.pdf — Fig. S1. Maximum Likelihood cladogram of the GENES-partitioned RAxML analysis of the molecular dataset. Numbers at internal branches are bootstrap values (BS) ≥ 50% in- ferred from 1,000 thorough bootstrap replicates. — DOI: 10.26049/ ASP77-1-2019-07/2

204