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

© Senckenberg Gesellschaft für Naturforschung, 2019.

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

Richard Mally *

, James E. Hayden

, Christoph Neinhuis

, Bjarte H. Jordal

& Matthias Nuss

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 Florida Department of Agriculture and Consumer Ser- vices, Division of Plant Industry, 1911 SW 34^th 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 moths). Morphological distinction of the two groups has been difficult 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 confirmed as ingroup of Spilomelinae, and Sufetula Walker, 1859 as a non-spilomeline. Within Spilomelinae, several well supported clades are obtained, for which we propose a first phylogeny-based tribal classification, 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., Spilomelini 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 Conchylodes Guenée, 1854, with Conchylodes graph­

ialis (Schaus, 1912) comb.n.; Conchylodes octonalis (Zeller, 1873) comb.n. (from Lygropia); Hyperectis Meyrick, 1904 syn.n. of Hydriris Meyrick, 1885, with Hydriris dioctias (Meyick, 1904) comb.n., and Hydriris apicalis (Hampson, 1912) comb.n.; Conogethes pandamalis (Walker, 1859) comb.n. (from Dichocrocis); Arthromastix pactolalis (Guenée, 1854) comb.n. (from Syllepte); Prophantis coenostolalis (Hampson, 1899) comb.n. (from Thliptoceras); Prophantis xanthomeralis (Hampson, 1918) comb.n. (from Thliptoceras); Prophantis longicornalis (Mabille, 1900) comb.n. (from Syngamia); Charitoprepes apicipicta (Inoue, 1963) comb.n. (from Heterocnephes); Prenesta rubrocinctalis (Guenée, 1854) comb.n. (from Glyphodes); Alytana calligrammalis (Mabille, 1879) comb.n. (from Analyta). 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 Odontiinae; Orthoraphis Hampson, 1896 from Spi- lomelinae to Lathrotelinae; Hydropionea Hampson, 1917, Plantegumia Amsel, 1956 and Munroe’s (1995) “undescribed genus ex Boeo­

tarcha Meyrick” are transferred from Spilomelinae to Glaphyriinae.

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

1. Introduction

Pyraustinae and Spilomelinae comprise over 5,200 de- scribed species worldwide, accounting for about one third of the species in Pyraloidea (n^uss et al. 2003 – 2019). It is estimated that about 50% of the pyraloid species are still undescribed on a global scale (M^unroe 1972a), and s^ut-

ton et al. (2015) estimated that in Southeast Asia 60% of the species remain to be described. The knowledge about eco logy and especially host plant associations of the lar- vae is rather comprehensive for the species occurring in Europe (e.g. H^asenfuss 1960; e^MMet 1988) and North America (e.g. M^unroe 1972a,b, 1974b, 1976a). A review of the known host plant data for the Oriental species has been given by r^obinson et al. (2001). In recent years, rear- ing efforts of Lepidoptera caterpillars like those in Papua New Guinea (M^iller et al. 2007) and Costa Rica (J^anzen

& H^allwacHs 2009) collected from the local flora have accumulated a large amount of host plant data for tropi- cal Spilomelinae. Altogether, spilomeline larvae feed on a large variety of angiosperms, with varying degrees of host specificity, and a few species feed on gymnosperms (e.g. i^noue & Y^aManaka 2006) and ferns (e.g. f^araH-

pour-H^agani et al. 2016). Larvae of Niphopyralis Hamp- son, 1893 are associated with weaver ants, living in their nests and feeding on ant larvae (r^oepke 1916; k^eMner 1923). Several species are known for their economic im- pact on crops, among them the corn borers of the genus Ostrinia Hübner, 1825 (e.g. n^afus & s^cHreiner 1991), the bean pod borer Maruca vitrata (Fabricius, 1787) (e.g.

s^HarMa 1998), the rice leafrollers of the genera Cnapha­

locrocis Lederer, 1863 and Marasmia Lederer, 1863 (e.g.

p^atHak & k^Han 1994) as well as the eggplant borers in the genus Leucinodes Guenée, 1854 (e.g. M^allY et al.

2015). Corn borers of the genus Ostrinia have become model systems in basic and applied research, like for population ecology, genetics and management as well as pheromone research (e^lswortH et al. 1989; b^urgio &

M^ani 1995; o^nstad & g^ould 1998; w^ang et al. 1998;

r^oelofs et al. 2002; l^assance 2010; f^uJi et al. 2011).

Though there has been continuous progress in the systematics of Spilomelinae and Pyraustinae, their cur- rent classification is still largely based on typological concepts. Spilomelinae and Pyraustinae are not easily distinguishable based on external features and therefore have long been considered as one taxon under the name Pyraustidae, together with distinct groups like Schoeno- biinae, Acentropinae, Scopariinae, Odontiinae and Gla- phy riinae (M^arion 1952). The distinction between Spi- lo melinae and Pyraustinae began to come into focus through analyses of genitalia by M^üller-r^utz (1929), p^ierce & M^etcalfe (1938) and M^arion (1952, 1954).

During the 1970s, the consensus was to classify Pyraus- tinae into Spilomelini and Pyraustini (M^unroe 1964, 1976a, 1995; M^unroe & s^olis 1998). In 1982, M^inet split Spilomelinae from Pyraustinae, regarding them only distantly related based on the lack of convincing synapomorphies. He considered the bilobed praecinc-

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 diagnostic features for Spilomelinae to be uniquely au- tapomorphic; instead, he diagnosed Spilomelinae by a combination of characters: chaetosemata absent, males without subcostal retinaculum, praecinctorium bilobed, tympanic frame protruding, spinulae distinctly tapered, male genitalia without well-developed gnathos, and fe- male genitalia without large rhombical signum. s^olis

& M^aes’ (2003) cladistic study based on morphological features of adults also implied that Pyraustinae and Spi- lomelinae are not closely related. In contrast, a phyloge- netic analysis of molecular data by r^egier et al. (2012) supported the monophyly of Pyraustinae + Spilomelinae;

the diversity of both groups, however, was poorly sam- pled, with only two species of Pyraustinae and three spe- cies of Spilomelinae included. Wurthiinae, characterised by a number of morphological adaptations to their ant association, was recovered as ingroup of Spilomelinae.

Recently, Lathrotelinae was revised and removed from Spilomelinae, comprising Diplopseustis Meyrick, 1884, Diplopseustoides Guillermet, 2013, Lathroteles J.F.G.

Clarke, 1971 and Sufetula Walker, 1859 (Minet 2015).

The classification of Spilomelinae is confusing. The subfamily includes 4,097 described species in 338 genera (n^uss et al. 2003 – 2019). Many genera contain only a few species, and 87 genera (26%) are monotypic. In contrast, 20 genera comprise more than 50 species, collectively encompassing 50% of the species. The most species-rich genera are Udea Guenée, 1845, Palpita Hübner, 1808, Glyphodes Guenée, 1854 and the heterogeneous genera Syllepte Hübner, 1823 and Lamprosema Hübner, 1823.

Pyraustinae comprises 1,239 described species in 174 genera, with 94 genera (52%) monotypic and only three genera with more than 50 species: Loxostege Hübner, 1825, Anania Hübner, 1823 and Pyrausta Schrank, 1802 (n^uss et al. 2003 – 2019). Tribes within Spilomelinae and Py rau stinae have been proposed for recognition in the past, but they usually served to segregate single genera with aberrant morphology, e.g. the long-legged, narrow- winged Lineodini Amsel, 1956, Nomophilini Kuznetzov

& Stekolnikov, 1979 and Hydririni Minet, 1982. There- fore, a comprehensive tribal classification has not been thoroughly accepted. M^unroe (1995) classified the Neo- tropical Spilomelinae into 15 genus groups plus many unplaced genera, but he did not provide diagnoses for these informal genus groups.

The natural relationships among some Spilomelinae genera have been investigated (s^utrisno 2002a,b, 2003, 2004, 2005, 2006; s^utrisno et al. 2006; M^allY & n^uss 2010; H^aines & r^ubinoff 2012), but a large-scale phy- logenetic analysis that takes the outstanding diversity of Spilomelinae and Pyraustinae into account and identifies main lineages and their phylogenetic relationships has not been published to date.

Our study provides the first phylogenetic intra-sub- family classification of Spilomelinae and Pyraustinae based on analysis of molecular, morphological and eco-

Table 1. Genetically investigated material with accession numbers. Second row of header states the primer pairs used to amplify the respective genetic marker. (NCBI) in columns Origin and Collection refers to sequences obtained from GenBank, and neither origin nor storing collection are known. DNA sampleGenusSpeciesTribusOriginCollectionCOI (1440bp)EF-1a (1071bp) HybLCO/HybNancyHybJerry/HybPatHybOscar-6143/Bosie-6144HybEF51.9/EFrcM4 MTD393Synaphepunctalis (Fabricius, 1775)(PYRALINAE)GermanyMTDJF497027MK459848 MM00051Pyralisfarinalis (Linnaeus, 1758)(PYRALINAE)(NCBI)(NCBI)GU828590GU828389GU828925GU829221 MM09194Schoenobiusgigantellus (Denis & Schiffermüller, 1775)(SCHOENOBIINAE)(NCBI)(NCBI)GU828842GU929806GU829143GU829411 MM11223Clepsicosmairidia Meyrick, 1888(ACENTROPINAE)(NCBI)(NCBI)GU828852GU929816GU829154GU829419 MM03362Crambusuliginosellus Zeller, 1850(CRAMBINAE)(NCBI)(NCBI)GU828691GU828487GU829014GU829302 MM04967Eudoniatruncicolella (Stainton, 1849)(SCOPARIINAE)(NCBI)(NCBI)GU828709GU828504GU829032GU829321 MTD370Midilaguianensis Munroe, 1970(MIDILINAE)French GuianaMTDMK459667MK459849 MTD1307Sufetuladiminutalis (Walker, 1866)(LATHROTELINAE)GermanyMTDMK459668MK459850 ZMBN094Aetholixcf. flavibasalis (Guenée, 1854)AgroteriniMalaysiaZMBNMK459669MK459851 MTD1016Agathodesdesignalis Guenée, 1854MargaroniiniPeruZSMMK459670MK459852 MTD1328 (WPH221)Agrioglyptaexcelsalis (Walker, 1866)MargaroniiniAustraliaUHIMJX017869JX017948MK459853 MTD488AAgroteranemoralis (Scopoli, 1863)AgroteriniGermanyMTDMK459671MK459854 MTD1354Anageshnacf. primordialis (Dyar, 1906)SteniiniBoliviaMTDMK506102MK459672MK459855 MTD798Antigastracatalaunalis (Duponchel, 1833)MargaroniiniMoroccoMTDMK459673MK459856 MTD1331Apilocrocisnovateutonialis Munroe, 1968WurthiiniPeruZSMMK506080MK459674MK459857 MTD668Aristebuleaprincipis Munroe & Mutuura, 1968WurthiiniChinaMTDJF852437MK459675MK459858 MTD797Arnianervosalis (Guenée, 1854)NomophiliniMoroccoMTDMK459676MK459859 MTD776Arthromastixlauralis (Walker, 1859)AsciodiniVenezuelaMHNGJF852400MK459677MK459860 MTD1061Arthromastixpactolalis (Guenée, 1854)AsciodiniFrench GuianaR. RougerieJN305177MK459678MK459861 MTD1325Asciodescf. gordialis Guenée, 1854AsciodiniBoliviaMTDMK506100MK459679MK459862 MTD1019Asturodesfimbriauralis (Guenée, 1854)MargaroniiniPeruZSMMK506101MK459680MK459863 MTD1347Ategumiaebulealis (Guenée, 1854)NomophiliniBoliviaMTDMK506088MK459681MK459864 MTD1329Azochiscf. rufidiscalis Hampson, 1904MargaroniiniPeruZSMMK506095MK459682MK459865 MTD882Bocchoriscf inspersalis (Zeller, 1852)NomophiliniSierra LeoneT. KarischMK459683MK459866 MTD1281Botyodesdiniasalis (Walker, 1859)MargaroniiniChinaMTDMK506074MK459684MK459867 MTD1319Cadarenapudoraria (Hübner, 1825)MargaroniiniCameroonA. ZwickMK459685MK459868 MTD826Cnaphalocrociscf. medinalis (Guenée, 1854)SpilomeliniPhilippinesMTDMK459686MK459869 MTD1041Conchylodeszebra (Sepp, 1850)UdeiniFrench GuianaMTDMK506103MK459687MK459870 ITBC058Conogethespandamalis (Walker, 1859)MargaroniiniMalaysiaZMBNMK459688MK459871 MTD649Cydalimaperspectalis (Walker, 1859)MargaroniiniChinaMTDJF852281MK459689MK459872 MTD1047Desmiacf. tages (Cramer, 1777)NomophiliniFrench GuianaMTDMK506091MK459690MK459873 MTD1323Diaphaniahyalinata (Linnaeus, 1767)MargaroniiniBoliviaMTDMK506110MK459691MK459874 MTD557Diasemiareticularis (Linnaeus, 1761)NomophiliniRomaniaMTDMK459692MK459875 MTD1357Diasemiopsisleodocusalis (Walker, 1859)NomophiliniBoliviaMTDMK506106MK459693MK459876 ZMBN097Dichocrociscf. zebralis (Moore, 1867)MargaroniiniMalaysiaZMBNMK459694MK459877 MTD868Dolicharthriapunctalis (Denis & Schiffermüller, 1775)SteniiniSpainMTDMK459695MK459878 MTD786Duponcheliafovealis Zeller, 1847SteniiniMoroccoMTDMK459696MK459879 MTD1316Eporidiadariusalis Walker, 1859SpilomeliniCameroonA. ZwickMK459697MK459880

DNA sampleGenusSpeciesTribusOriginCollectionCOI (1440bp)EF-1a (1071bp) HybLCO/HybNancyHybJerry/HybPatHybOscar-6143/Bosie-6144HybEF51.9/EFrcM4 MTD1321Eurrhyparodescf. lygdamis Druce, 1902HerpetogrammatiniBoliviaMTDMK506099MK459698MK459881 MTD1566Filodessp.MargaroniiniAngolaMTDMK459699MK459882 MTD1318Ghesquierellanacf. hirtusalis (Walker, 1859)MargaroniiniCameroonA. ZwickMK459700MK459883 MTD1285Glyphodessibillalis Walker, 1859MargaroniiniPeruZSMMK459701MK459884 MTD820Glyphodescf. stolalis Guenée, 1854MargaroniiniPhilippinesMTDMK459702MK459885 MTD1283Gonocaustasp.HydririniBoliviaMTDMK506090MK459703MK459886 MTD808Haritalodesderogata (Fabricius, 1775)AgroteriniPhilippinesMTDMK459704MK459887 MTD994Herpetogrammaphaeopteralis (Guenée, 1854)HerpetogrammatiniPeruZSMMK506075MK459705MK459888 MTD1337Hileithiacf. obliqualis (Schaus, 1912)HerpetogrammatiniPeruZSMMK506096MK459706MK459889 MTD1282Hodebertiatestalis (Fabricius, 1794)MargaroniiniYemenMTDMK459707MK459890 MTD1565Hydririsornatalis (Duponchel, 1832)HydririniAngolaMTDMK459708MK459891 MTD1004Hymeniaperspectalis (Hübner, 1796)HymeniiniPeruZSMMK506081MK459709MK459892 MTD1043Lamprosemacf. dorisalis (Walker, 1859)HydririniFrench GuianaMTDMK506082MK459710MK459893 MTD1562Leucinodesafricensis Mally et al., 2015LineodiniAngolaMTDLN624711MK459711MK459894 MTD1349Leucochromacorope (Stoll in Cramer & Stoll, 1781)MargaroniiniBoliviaMTDMK506087MK459712MK459895 MTD1251Lineodesvulnifica Dyar, 1913LineodiniBoliviaMTDMK506112MK459713MK459896 MTD1284Liopasiaandrealis Dognin, 1910MargaroniiniBoliviaMTDMK506105MK459714MK459897 WPH197Marasmiapoeyalis (Boisduval, 1833)Spilomelini(NCBI)(NCBI)JX017856JX017856JX017933— WPH115Marasmiatrapezalis (Guenée, 1854)Spilomelini(NCBI)(NCBI)JX017849JX017849JX017926— MTD1341Marucavitrata (Fabricius, 1787)MargaroniiniBoliviaMTDMK506085MK459715MK459898 MTD364Mecynalutealis (Duponchel, 1833)NomophiliniItalyTLMFJF497031JF497031MK459899 MTD1340Megastescf. pusialis Snellen, 1875MargaroniiniBoliviaMTDMK506098MK459716MK459900 MTD787Metasiasuppandalis (Hübner, 1823)SteniiniMoroccoMTDMK459717MK459901 ZMBN104Nacoleiainsolitalis (Walker, 1862)MargaroniiniMalaysiaZMBNMK459718MK459902 ZMBN103Neoanalthescf. pseudocontortalis Yamanaka & Kirpichnikova, 1993AgroteriniMalaysiaZMBNMK459719MK459903 MTD1046Neoleucinodesdissolvens (Dyar, 1914)LineodiniFrench GuianaMTDMK506093MK459720MK459904 MTD152Niphopyralischionesis Hampson, 1919WurthiiniAustraliaANICMK459721MK459905 MTD782Nomophilanoctuella (Denis & Schiffermüller, 1775)NomophiliniMoroccoMTDMK459722MK459906 MTD1483Obtusipalpispardalis Hampson, 1896MargaroniiniAngolaMTDMK459723MK459907 SDA008AOmiodescontinuatalis (Wallengren, 1860)MargaroniiniHawaiiUHIMMK459724MK459908 WPH252BOmiodeshumeralis Guenée, 1854MargaroniiniCosta RicaUHIMJX017886JX017886JX017965MK459909 ZMBN097Dichocrociscf. zebralis (Moore, 1867)MargaroniiniMalaysiaZMBNMK459694MK459877 MTD784Palpitavitrealis (Rossi, 1794)MargaroniiniMoroccoMTDMK459725MK459910 MM00325Pataniaruralis (Scopoli, 1763)Agroterini(NCBI)(NCBI)GU828634GU828432GU828968GU829254 MTD1018Pataniacf. silicalis (Guenée, 1854)AgroteriniPeruZSMMK506097MK459726MK459911 MTD1324Phostriacf. tedea (Stoll in Cramer & Stoll, 1780)AgroteriniBoliviaMTDMK506092MK459727MK459912 MTD1033Prenestacf. iphiclalis (Walker, 1859)MargaroniiniFrench GuianaMTDMK506083MK459728—MK459913 MTD1342Prenestacf. rubrocinctalis (Guenée, 1854)MargaroniiniBoliviaMTDMK506078MK459729—MK459914

Table 1 continued.

Table 1 continued. DNA sampleGenusSpeciesTribusOriginCollectionCOI (1440bp)EF-1a (1071bp) HybLCO/HybNancyHybJerry/HybPatHybOscar-6143/Bosie-6144HybEF51.9/EFrcM4 MTD1015Prenestascyllalis (Walker, 1859)MargaroniiniPeruZSMMK506094MK459730MK459915 WPH188Prophantiscf. androstigmata (Hampson, 1918)Trichaeini(NCBI)(NCBI)JX017853JX017853JX017930— ZMBN017Prophantisxanthomeralis (Hampson, 1918)TrichaeiniAngolaMTDMK459731MK459916 MTD650Pycnarmonpantherata (Butler, 1878)AgroteriniChinaMTDMK459732MK459917 MTD774Rhectosemiamultifarialis Lederer, 1863LineodiniVenezuelaMHNGJF852398MK459733MK459918 MTD874Rhimphaleacf. astrigalis Hampson, 1899MargaroniiniPhilippinesMTDMK459734MK459919 ZMBN011Salbiahaemorrhoidalis (Guenée, 1854)SpilomeliniBoliviaMTDMK506086MK459735MK459920 MTD1358Sameacf. multiplicalis (Guenée, 1854)NomophiliniBoliviaMTDMK506079MK459736MK459921 MTD1235Sameaecclesialis Guenée, 1854NomophiliniUSAFMNHMK459737MK459922 MTD872Sigaliris (Cramer, 1775)SpilomeliniFrench GuianaMHNGMK459738MK459923 MTD1248Spilomelaperspicata (Fabricius, 1787)SpilomeliniPeruZSMMK506084MK459739MK459924 MTD783Spoladearecurvalis (Fabricius, 1775)HymeniiniMoroccoMTDMK459740MK459925 MTD1320Syllepismarialis Poey, 1832HydririniBoliviaMTDLR135741LR135741MK459926 MTD1017Syngamiaflorella (Stoll in Cramer & Stoll, 1781)SpilomeliniPeruZSMMK506076MK459741MK459927 MTD1315Terastiameticulosalis Guenée, 1854MargaroniiniPeruZSMMK506111MK459742MK459928 MTD1247Trichaeapilicornis Herrich-Schäffer, 1866TrichaeiniPeruZSMMK506089MK459743MK459929 MTD870Udeaferrugalis (Hübner, 1796)UdeiniMoroccoMTDJF852252MK459744MK459930 MTD956Udeawashingtonalis (Grote, 1882)UdeiniCanadaMTDMK459745MK459931 MTD276, MTD357Udeoidesmuscosalis (Hampson, 1913)UdeiniKenyaMTDJF497033JF497033MK459932 MTD1467Zebroniaphenice (Stoll in Cramer & Stoll, 1782)MargaroniiniAngolaMTDMK459746MK459933 MTD1338Achyracf. rantalis (Guenée, 1854)PyraustiniBoliviaMTDMK506109MK459747MK459934 MM01851Ananiahortulata (Linnaeus, 1758)Pyraustini(NCBI)(NCBI)GU828675GU828472GU829003GU829287 MTD553Ananiaverbascalis (Denis & Schiffermüller, 1775)PyraustiniRomaniaMTDMK459748MK459935 MTD1484Cryptosaracaritalis (Walker, 1859)PortentomorphiniAngolaMTDMK459749MK459936 MTD1558Euclastagigantalis Viette, 1957EuclastiniKenyaNHMOMK459750MK459937 MTD1466Euclastasplendidalis (Herrich-Schäffer, 1848)EuclastiniBulgariaS. BeshkovMK459751MK459938 MTD1327 (WPH215)Hyalobathracrenulata Sutrisno & Horak, 2003PortentomorphiniAustraliaUHIMJX017826JX017826JX017943MK459939 MTD1350Hyaloristacf. taeniolalis (Guenée, 1854)PyraustiniBoliviaMTDMK506108MK459752MK459940 MTD605Loxostegeaeruginalis (Hübner, 1796)PyraustiniMacedoniaMTDMK459753MK459941 MTD1343Oenobotyssp.PyraustiniBoliviaMTDMK506104MK459754MK459942 MTD388Ostrinianubilalis (Hübner, 1796)PyraustiniGermanyMTDMK459755MK459943 ZMBN096Pagydasalvalis Walker, 1859PyraustiniMalaysiaZMBNMK459756MK459944 MTD906Paracorsiarepandalis (Denis & Schiffermüller, 1775)PyraustiniKyrgyzstanN. PöllMK459757MK459945 MTD1322Portentomorphaxanthialis (Guenée, 1854)PortentomorphiniBoliviaMTDMK506077MK459758MK459946 MTD477Psammotispulveralis (Hübner, 1796)PyraustiniGermanyMTDMK459759MK459947 MTD1344Pseudopyraustacf. minima (Hedemann, 1894)PyraustiniBoliviaMTDMK506107MK459760MK459948

Table 1 continued. DNA sampleGenusSpeciesTribusOriginCollectionCOI (1440bp)EF-1a (1071bp) HybLCO/HybNancyHybJerry/HybPatHybOscar-6143/Bosie-6144HybEF51.9/EFrcM4 MTD953Pyraustapurpuralis (Linnaeus, 1758)PyraustiniGermanyMTDMK459761MK459949 MTD560Sitochroaverticalis (Linnaeus, 1758)PyraustiniRomaniaMTDMK459762MK459950 MTD1326 (WPH209)Tetridiacaletoralis (Walker, 1859)TetridiiniAustraliaUHIMJX017861JX017861JX017938MK459951 WPH054Uresiphitagilvata (Fabricius, 1794)unplacedHawaiiUHIMJX017825JX017825JX017919— DNA sampleGenusSpeciesTribusOriginCollectionCAD (825bp)GAPDH (654bp)IDH (657bp)RpS5 (576bp) HybCAD743f/ HybCAD1028rHybFrigga/ HybBurreHybIDHdeg27F/ HybIDHdegRHybRpS5f/ HybRpS5r MTD393Synaphepunctalis (Fabricius, 1775)(PYRALINAE)GermanyMTDMK459763MK460136MK459952MK460052 MM00051Pyralisfarinalis (Linnaeus, 1758)(PYRALINAE)(NCBI)(NCBI)GU828092GU829747GU829979GU830604 MM09194Schoenobiusgigantellus (Denis & Schiffermüller, 1775)(SCHOENOBIINAE)(NCBI)(NCBI)GU828306GU829903GU830222GU830790 MM11223Clepsicosmairidia Meyrick, 1888(ACENTROPINAE)(NCBI)(NCBI)GU828315GU829906GU830230GU830800 MM03362Crambusuliginosellus Zeller, 1850(CRAMBINAE)(NCBI)(NCBI)GU828182GU829811GU830078GU830681 MM04967Eudoniatruncicolella (Stainton, 1849)(SCOPARIINAE)(NCBI)(NCBI)GU828197GU829823GU830095GU830697 MTD370Midilaguianensis Munroe, 1970(MIDILINAE)French GuianaMTDMK459764MK460137MK459953MK460053 MTD1307Sufetuladiminutalis (Walker, 1866)(LATHROTELINAE)GermanyMTDMK459765MK460138MK459954MK460054 ZMBN094Aetholixcf. flavibasalis (Guenée, 1854)AgroteriniMalaysiaZMBNMK459766MK460139MK459955MK460055 MTD1016Agathodesdesignalis Guenée, 1854MargaroniiniPeruZSMMK459767—MK459956MK460056 MTD1328 (WPH221)Agrioglyptaexcelsalis (Walker, 1866)MargaroniiniAustraliaUHIMJX017793—MK459957JX018024 MTD488AAgroteranemoralis (Scopoli, 1863)AgroteriniGermanyMTDMK459768MK460140MK459958MK460057 MTD1354Anageshnacf. primordialis (Dyar, 1906)SteniiniBoliviaMTDMK459769MK460141MK459959MK460058 MTD798Antigastracatalaunalis (Duponchel, 1833)MargaroniiniMoroccoMTDMK459770—MK459960MK460059 MTD1331Apilocrocisnovateutonialis Munroe, 1968WurthiiniPeruZSMMK459771—MK459961— MTD668Aristebuleaprincipis Munroe & Mutuura, 1968WurthiiniChinaMTDMK459772MK460142MK459962— MTD797Arnianervosalis (Guenée, 1854)NomophiliniMoroccoMTDMK459773——MK460060 MTD776Arthromastixlauralis (Walker, 1859)AsciodiniVenezuelaMHNG—MK460143MK459963MK460061 MTD1061Arthromastixpactolalis (Guenée, 1854)AsciodiniFrench GuianaR. RougerieMK459774MK460144MK459964MK460062 MTD1325Asciodescf. gordialis Guenée, 1854AsciodiniBoliviaMTDMK459775MK460145MK459965MK460063 MTD1019Asturodesfimbriauralis (Guenée, 1854)MargaroniiniPeruZSMMK459776—MK459966MK460064 MTD1347Ategumiaebulealis (Guenée, 1854)NomophiliniBoliviaMTDMK459777—MK459967MK460065 MTD1329Azochiscf. rufidiscalis Hampson, 1904MargaroniiniPeruZSMMK459778MK460146MK459968MK460066 MTD882Bocchoriscf. inspersalis (Zeller, 1852)NomophiliniSierra LeoneT. Karisch—MK460147—MK460067 MTD1281Botyodesdiniasalis (Walker, 1859)MargaroniiniChinaMTD——MK459969MK460068 MTD1319Cadarenapudoraria (Hübner, 1825)MargaroniiniCameroonA. ZwickMK459779MK460148MK459970MK460069 MTD826Cnaphalocrociscf. medinalis (Guenée, 1854)SpilomeliniPhilippinesMTDMK459780—MK459971MK460070 MTD1041Conchylodeszebra (Sepp, 1850)UdeiniFrench GuianaMTDMK459781MK460149MK459972— ITBC058Conogethespandamalis (Walker, 1859)MargaroniiniMalaysiaZMBNMK459782MK460150MK459973MK460071

Table 1 continued. DNA sampleGenusSpeciesTribusOriginCollectionCAD (825bp)GAPDH (654bp)IDH (657bp)RpS5 (576bp) HybCAD743f/ HybCAD1028rHybFrigga/ HybBurreHybIDHdeg27F/ HybIDHdegRHybRpS5f/ HybRpS5r MTD649Cydalimaperspectalis (Walker, 1859)MargaroniiniChinaMTD—MK460151MK459974— MTD1047Desmiacf. tages (Cramer, 1777)NomophiliniFrench GuianaMTDMK459783MK460152MK459975MK460072 MTD1323Diaphaniahyalinata (Linnaeus, 1767)MargaroniiniBoliviaMTDMK459784MK460153MK459976MK460073 MTD557Diasemiareticularis (Linnaeus, 1761)NomophiliniRomaniaMTDMK459785MK460154MK459977MK460074 MTD1357Diasemiopsisleodocusalis (Walker, 1859)NomophiliniBoliviaMTDMK459786MK460155MK459978MK460075 ZMBN097Dichocrociscf. zebralis (Moore, 1867)MargaroniiniMalaysiaZMBNMK459787MK460156MK459979MK460076 MTD868Dolicharthriapunctalis (Denis & Schiffermüller, 1775)SteniiniSpainMTDMK459788—MK459980MK460077 MTD786Duponcheliafovealis Zeller, 1847SteniiniMoroccoMTDMK459789MK460157MK459981MK460078 MTD1316Eporidiadariusalis Walker, 1859SpilomeliniCameroonA. ZwickMK459790—MK459982MK460079 MTD1321Eurrhyparodescf. lygdamis Druce, 1902HerpetogrammatiniBoliviaMTDMK459791MK460158MK459983MK460080 MTD1566Filodessp.MargaroniiniAngolaMTDMK459792—MK459984MK460081 MTD1318Ghesquierellanacf. hirtusalis (Walker, 1859)MargaroniiniCameroonA. ZwickMK459793MK460159MK459985MK460082 MTD1285Glyphodessibillalis Walker, 1859MargaroniiniPeruZSMMK459794MK460160MK459986MK460083 MTD820Glyphodescf. stolalis Guenée, 1854MargaroniiniPhilippinesMTDMK459795MK460161MK459987MK460084 MTD1283Gonocaustasp.HydririniBoliviaMTDMK459796MK460162MK459988MK460085 MTD808Haritalodesderogata (Fabricius, 1775)AgroteriniPhilippinesMTDMK459797MK460163MK459989— MTD994Herpetogrammaphaeopteralis (Guenée, 1854)HerpetogrammatiniPeruZSMMK459798—MK459990MK460086 MTD1337Hileithiacf. obliqualis (Schaus, 1912)HerpetogrammatiniPeruZSMMK459799MK460164MK459991MK460087 MTD1282Hodebertiatestalis (Fabricius, 1794)MargaroniiniYemenMTDMK459800MK460165MK459992MK460088 MTD1565Hydririsornatalis (Duponchel, 1832)HydririniAngolaMTDMK459801MK460166—— MTD1004Hymeniaperspectalis (Hübner, 1796)HymeniiniPeruZSMMK459802MK460167MK459993MK460089 MTD1043Lamprosemacf. dorisalis (Walker, 1859)HydririniFrench GuianaMTDMK459803MK460168MK459994MK460090 MTD1562Leucinodesafricensis Mally et al., 2015LineodiniAngolaMTDMK459804—MK459995— MTD1349Leucochromacorope (Stoll in Cramer & Stoll, 1781)MargaroniiniBoliviaMTDMK459805MK460169MK459996MK460091 MTD1251Lineodesvulnifica Dyar, 1913LineodiniBoliviaMTDMK459806MK460170MK459997MK460092 MTD1284Liopasiaandrealis Dognin, 1910MargaroniiniBoliviaMTDMK459807MK460171MK459998MK460093 WPH197Marasmiapoeyalis (Boisduval, 1833)Spilomelini(NCBI)(NCBI)JX017781——JX018009 WPH115Marasmiatrapezalis (Guenée, 1854)Spilomelini(NCBI)(NCBI)JX017777——JX018002 MTD1341Marucavitrata (Fabricius, 1787)MargaroniiniBoliviaMTDMK459808—MK459999MK460094 MTD364Mecynalutealis (Duponchel, 1833)NomophiliniItalyTLMF—MK460172MK460000MK460095 MTD1340Megastescf. pusialis Snellen, 1875MargaroniiniBoliviaMTDMK459809MK460173MK460001MK460096 MTD787Metasiasuppandalis (Hübner, 1823)SteniiniMoroccoMTDMK459810MK460174MK460002MK460097 ZMBN104Nacoleiainsolitalis (Walker, 1862)MargaroniiniMalaysiaZMBNMK459811—MK460003— ZMBN103Neoanalthescf. pseudocontortalis Yamanaka & Kirpichnikova, 1993AgroteriniMalaysiaZMBN—MK460175MK460004MK460098 MTD1046Neoleucinodesdissolvens (Dyar, 1914)LineodiniFrench GuianaMTDMK459812MK460176MK460005— MTD152Niphopyralischionesis Hampson, 1919WurthiiniAustraliaANICMK459813—MK460006MK460099 MTD782Nomophilanoctuella (Denis & Schiffermüller, 1775)NomophiliniMoroccoMTDMK459814MK460177MK460007MK460100 MTD1483Obtusipalpispardalis Hampson, 1896MargaroniiniAngolaMTDMK459815MK460178MK460008MK460101