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

May 2019
Arthropod Systematics and Phylogeny 77(1):141-204

May 2019
77(1):141-204

DOI:10.26049/ASP77-1-2019-07

Authors:

Richard Mally

Czech University of Life Sciences Prague

James E. Hayden

Florida Department of Agriculture and Consumer Services

Bjarte H Jordal

University of Bergen

Show all 5 authorsHide

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 examine 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 graphialis (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 Spilomelinae to Lathrotelinae; Hydropionea Hampson, 1917, Plantegumia Amsel, 1956 and Munroe’s (1995) “undescribed genus ex Boeotarcha Meyrick” are transferred from Spilomelinae to Glaphyriinae.

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 represents 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.

…

Strict consensus of three parsimony cladograms of 35,100 steps, with morphological characters mapped with slow optimization. -Symbols: • unique apomorphies; ○ homoplastic apomorphies.

…

Continuation.

…

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.

…

Lineodini. A: ventral view of wings on frenulum bristle of female Leucinodes orbonalis. B: head of Le. orbonalis, male (left) and female (right) (modified from Figs. 11 -12 of MallY et al. 2015). C: adult male of Lineodes vulnifica. D: adult female of Le. Africensis. E: male genitalia of Li. vulnifica. 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.

…

Figures - uploaded by Richard Mally

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141

ISSN 1863-7221 (print)

eISSN 1864-8312 (online)

DOI: 10.26049/ASP77-1-2019-07

(1): 141 – 204

2019

The phylogenetic systematics of Spilomelinae and

Pyraustinae (Lepidoptera: Pyraloidea: Crambidae)

inferred from DNA and morphology

R M *, 1, J E. H 2, C N 3,

B H. J 1 & M N 4

University Museum of Bergen, Natural History Collections, Realfagbygget, Allégaten 41, 5007 Bergen, Norway; Richard Mally [richard.

mally@uib.no, spilomallynae@gmail.com], Bjarte H. Jordal [bjarte.jordal@uib.no] —

Florida Department of Agriculture and Consumer Ser-

vices, Division of Plant Industry, 1911 SW 34

Street, Gainesville, FL 32608 USA; James E. Hayden [james.hayden@freshfromflorida.com] —

Technische Universität Dresden, Institut für Botanik, 01062 Dresden, Germany; Christoph Neinhuis [christoph.neinhuis@tu-dresden.de]

—

Senckenberg Naturhistorische Sammlungen Dresden, Museum für Tierkunde, Königsbrücker Landstraße 159, 01109 Dresden, Germany;

Matthias Nuss [matthias.nuss@senckenberg.de] — * 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 difcult 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 conrmed 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

rst phylogeny-based tribal classication, 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 classication, morphology.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

142

1. Introduction

Pyraustinae and Spilomelinae comprise over 5,200 de-

scribed species worldwide, accounting for about one third

of the species in Pyraloidea (nuss et al. 2003 – 2019). It is

estimated that about 50% of the pyraloid species are still

undescribed on a global scale (Munroe 1972a), and sut-

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. Hasenfuss 1960; eMMet 1988) and North

America (e.g. Munroe 1972a,b, 1974b, 1976a). A review

of the known host plant data for the Oriental species has

been given by robinson et al. (2001). In recent years, rear-

ing efforts of Lepidoptera caterpillars like those in Papua

New Guinea (Miller et al. 2007) and Costa Rica (Janzen

& HallwacHs 2009) collected from the local ora 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 specicity, and a few species feed on gymnosperms

(e.g. inoue & YaManaka 2006) and ferns (e.g. faraH-

pour-Hagani et al. 2016). Larvae of Niphopyralis Hamp-

son, 1893 are associated with weaver ants, living in their

nests and feeding on ant larvae (roepke 1916; keMner

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. nafus & scHreiner 1991),

the bean pod borer Maruca vitrata (Fabricius, 1787) (e.g.

sHarMa 1998), the rice leafrollers of the genera Cnapha-

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

patHak & kHan 1994) as well as the eggplant borers in

the genus Leucinodes Guenée, 1854 (e.g. MallY 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 (elswortH et al. 1989; burgio &

Mani 1995; onstad & gould 1998; wang et al. 1998;

roelofs et al. 2002; lassance 2010; fuJi et al. 2011).

Though there has been continuous progress in the

systematics of Spilomelinae and Pyraustinae, their cur-

rent classication 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 (Marion 1952). The distinction between Spi-

lo melinae and Pyraustinae began to come into focus

through analyses of genitalia by Müller-rutz (1929),

pierce & Metcalfe (1938) and Marion (1952, 1954).

During the 1970s, the consensus was to classify Pyraus-

tinae into Spilomelini and Pyraustini (Munroe 1964,

1976a, 1995; Munroe & solis 1998). In 1982, Minet

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. solis

& Maes’ (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 regier 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 classication of Spilomelinae is confusing. The

subfamily includes 4,097 described species in 338 genera

(nuss 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

(nuss 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 classication has not been

thoroughly accepted. Munroe (1995) classied 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 (sutrisno 2002a,b, 2003,

2004, 2005, 2006; sutrisno et al. 2006; MallY & nuss

2010; Haines & rubinoff 2012), but a large-scale phy-

logenetic analysis that takes the outstanding diversity of

Spilomelinae and Pyraustinae into account and identies

main lineages and their phylogenetic relationships has

not been published to date.

Our study provides the rst phylogenetic intra-sub-

family classication of Spilomelinae and Pyraustinae

based on analysis of molecular, morphological and eco-

143

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

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 sample Genus Species Tribus Origin Collection COI (1440bp) EF-1a (1071bp)

HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4

MTD393 Synaphe punctalis (Fabricius, 1775) (PYRALINAE) Germany MTD JF497027 MK459848

MM00051 Pyralis farinalis (Linnaeus, 1758) (PYRALINAE) (NCBI) (NCBI) GU828590 GU828389 GU828925 GU829221

MM09194 Schoenobius gigantellus (Denis & Schiermüller, 1775) (SCHOENOBIINAE) (NCBI) (NCBI) GU828842 GU929806 GU829143 GU829411

MM11223 Clepsicosma iridia Meyrick, 1888 (ACENTROPINAE) (NCBI) (NCBI) GU828852 GU929816 GU829154 GU829419

MM03362 Crambus uliginosellus Zeller, 1850 (CRAMBINAE) (NCBI) (NCBI) GU828691 GU828487 GU829014 GU829302

MM04967 Eudonia truncicolella (Stainton, 1849) (SCOPARIINAE) (NCBI) (NCBI) GU828709 GU828504 GU829032 GU829321

MTD370 Midila guianensis Munroe, 1970 (MIDILINAE) French Guiana MTD MK459667 MK459849

MTD1307 Sufetula diminutalis (Walker, 1866) (LATHROTELINAE) Germany MTD MK459668 MK459850

ZMBN094 Aetholix cf. ﬂavibasalis (Guenée, 1854) Agroterini Malaysia ZMBN MK459669 MK459851

MTD1016 Agathodes designalis Guenée, 1854 Margaroniini Peru ZSM MK459670 MK459852

MTD1328

(WPH221) Agrioglypta excelsalis (Walker, 1866) Margaroniini Australia UHIM JX017869 JX017948 MK459853

MTD488A Agrotera nemoralis (Scopoli, 1863) Agroterini Germany MTD MK459671 MK459854

MTD1354 Anageshna cf. primordialis (Dyar, 1906) Steniini Bolivia MTD MK506102 MK459672 MK459855

MTD798 Antigastra catalaunalis (Duponchel, 1833) Margaroniini Morocco MTD MK459673 MK459856

MTD1331 Apilocrocis novateutonialis Munroe, 1968 Wurthiini Peru ZSM MK506080 MK459674 MK459857

MTD668 Aristebulea principis Munroe & Mutuura, 1968 Wurthiini China MTD JF852437 MK459675 MK459858

MTD797 Arnia nervosalis (Guenée, 1854) Nomophilini Morocco MTD MK459676 MK459859

MTD776 Arthromastix lauralis (Walker, 1859) Asciodini Venezuela MHNG JF852400 MK459677 MK459860

MTD1061 Arthromastix pactolalis (Guenée, 1854) Asciodini French Guiana R. Rougerie JN305177 MK459678 MK459861

MTD1325 Asciodes cf. gordialis Guenée, 1854 Asciodini Bolivia MTD MK506100 MK459679 MK459862

MTD1019 Asturodes ﬁmbriauralis (Guenée, 1854) Margaroniini Peru ZSM MK506101 MK459680 MK459863

MTD1347 Ategumia ebulealis (Guenée, 1854) Nomophilini Bolivia MTD MK506088 MK459681 MK459864

MTD1329 Azochis cf. ruﬁdiscalis Hampson, 1904 Margaroniini Peru ZSM MK506095 MK459682 MK459865

MTD882 Bocchoris cf inspersalis (Zeller, 1852) Nomophilini Sierra Leone T. Karisch MK459683 MK459866

MTD1281 Botyodes diniasalis (Walker, 1859) Margaroniini China MTD MK506074 MK459684 MK459867

MTD1319 Cadarena pudoraria (Hübner, 1825) Margaroniini Cameroon A. Zwick MK459685 MK459868

MTD826 Cnaphalocrocis cf. medinalis (Guenée, 1854) Spilomelini Philippines MTD MK459686 MK459869

MTD1041 Conchylodes zebra (Sepp, 1850) Udeini French Guiana MTD MK506103 MK459687 MK459870

ITBC058 Conogethes pandamalis (Walker, 1859) Margaroniini Malaysia ZMBN MK459688 MK459871

MTD649 Cydalima perspectalis (Walker, 1859) Margaroniini China MTD JF852281 MK459689 MK459872

MTD1047 Desmia cf. tages (Cramer, 1777) Nomophilini French Guiana MTD MK506091 MK459690 MK459873

MTD1323 Diaphania hyalinata (Linnaeus, 1767) Margaroniini Bolivia MTD MK506110 MK459691 MK459874

MTD557 Diasemia reticularis (Linnaeus, 1761) Nomophilini Romania MTD MK459692 MK459875

MTD1357 Diasemiopsis leodocusalis (Walker, 1859) Nomophilini Bolivia MTD MK506106 MK459693 MK459876

ZMBN097 Dichocrocis cf. zebralis (Moore, 1867) Margaroniini Malaysia ZMBN MK459694 MK459877

MTD868 Dolicharthria punctalis (Denis & Schiermüller, 1775) Steniini Spain MTD MK459695 MK459878

MTD786 Duponchelia fovealis Zeller, 1847 Steniini Morocco MTD MK459696 MK459879

MTD1316 Eporidia dariusalis Walker, 1859 Spilomelini Cameroon A. Zwick MK459697 MK459880

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

144

DNA sample Genus Species Tribus Origin Collection COI (1440bp) EF-1a (1071bp)

HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4

MTD1321 Eurrhyparodes cf. lygdamis Druce, 1902 Herpetogrammatini Bolivia MTD MK506099 MK459698 MK459881

MTD1566 Filodes sp. Margaroniini Angola MTD MK459699 MK459882

MTD1318 Ghesquierellana cf. hirtusalis (Walker, 1859) Margaroniini Cameroon A. Zwick MK459700 MK459883

MTD1285 Glyphodes sibillalis Walker, 1859 Margaroniini Peru ZSM MK459701 MK459884

MTD820 Glyphodes cf. stolalis Guenée, 1854 Margaroniini Philippines MTD MK459702 MK459885

MTD1283 Gonocausta sp. Hydririni Bolivia MTD MK506090 MK459703 MK459886

MTD808 Haritalodes derogata (Fabricius, 1775) Agroterini Philippines MTD MK459704 MK459887

MTD994 Herpetogramma phaeopteralis (Guenée, 1854) Herpetogrammatini Peru ZSM MK506075 MK459705 MK459888

MTD1337 Hileithia cf. obliqualis (Schaus, 1912) Herpetogrammatini Peru ZSM MK506096 MK459706 MK459889

MTD1282 Hodebertia testalis (Fabricius, 1794) Margaroniini Yemen MTD MK459707 MK459890

MTD1565 Hydriris ornatalis (Duponchel, 1832) Hydririni Angola MTD MK459708 MK459891

MTD1004 Hymenia perspectalis (Hübner, 1796) Hymeniini Peru ZSM MK506081 MK459709 MK459892

MTD1043 Lamprosema cf. dorisalis (Walker, 1859) Hydririni French Guiana MTD MK506082 MK459710 MK459893

MTD1562 Leucinodes africensis Mally et al., 2015 Lineodini Angola MTD LN624711 MK459711 MK459894

MTD1349 Leucochroma corope (Stoll in Cramer & Stoll, 1781) Margaroniini Bolivia MTD MK506087 MK459712 MK459895

MTD1251 Lineodes vulniﬁca Dyar, 1913 Lineodini Bolivia MTD MK506112 MK459713 MK459896

MTD1284 Liopasia andrealis Dognin, 1910 Margaroniini Bolivia MTD MK506105 MK459714 MK459897

WPH197 Marasmia poeyalis (Boisduval, 1833) Spilomelini (NCBI) (NCBI) JX017856 JX017856 JX017933 —

WPH115 Marasmia trapezalis (Guenée, 1854) Spilomelini (NCBI) (NCBI) JX017849 JX017849 JX017926 —

MTD1341 Maruca vitrata (Fabricius, 1787) Margaroniini Bolivia MTD MK506085 MK459715 MK459898

MTD364 Mecyna lutealis (Duponchel, 1833) Nomophilini Italy TLMF JF497031 JF497031 MK459899

MTD1340 Megastes cf. pusialis Snellen, 1875 Margaroniini Bolivia MTD MK506098 MK459716 MK459900

MTD787 Metasia suppandalis (Hübner, 1823) Steniini Morocco MTD MK459717 MK459901

ZMBN104 Nacoleia insolitalis (Walker, 1862) Margaroniini Malaysia ZMBN MK459718 MK459902

ZMBN103 Neoanalthes cf. pseudocontortalis Yamanaka & Kirpichnikova, 1993 Agroterini Malaysia ZMBN MK459719 MK459903

MTD1046 Neoleucinodes dissolvens (Dyar, 1914) Lineodini French Guiana MTD MK506093 MK459720 MK459904

MTD152 Niphopyralis chionesis Hampson, 1919 Wurthiini Australia ANIC MK459721 MK459905

MTD782 Nomophila noctuella (Denis & Schiermüller, 1775) Nomophilini Morocco MTD MK459722 MK459906

MTD1483 Obtusipalpis pardalis Hampson, 1896 Margaroniini Angola MTD MK459723 MK459907

SDA008A Omiodes continuatalis (Wallengren, 1860) Margaroniini Hawaii UHIM MK459724 MK459908

WPH252B Omiodes humeralis Guenée, 1854 Margaroniini Costa Rica UHIM JX017886 JX017886 JX017965 MK459909

ZMBN097 Dichocrocis cf. zebralis (Moore, 1867) Margaroniini Malaysia ZMBN MK459694 MK459877

MTD784 Palpita vitrealis (Rossi, 1794) Margaroniini Morocco MTD MK459725 MK459910

MM00325 Patania ruralis (Scopoli, 1763) Agroterini (NCBI) (NCBI) GU828634 GU828432 GU828968 GU829254

MTD1018 Patania cf. silicalis (Guenée, 1854) Agroterini Peru ZSM MK506097 MK459726 MK459911

MTD1324 Phostria cf. tedea (Stoll in Cramer & Stoll, 1780) Agroterini Bolivia MTD MK506092 MK459727 MK459912

MTD1033 Prenesta cf. iphiclalis (Walker, 1859) Margaroniini French Guiana MTD MK506083 MK459728 — MK459913

MTD1342 Prenesta cf. rubrocinctalis (Guenée, 1854) Margaroniini Bolivia MTD MK506078 MK459729 — MK459914

Table 1 continued.

145

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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Table 1 continued.

DNA sample Genus Species Tribus Origin Collection COI (1440bp) EF-1a (1071bp)

HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4

MTD1015 Prenesta scyllalis (Walker, 1859) Margaroniini Peru ZSM MK506094 MK459730 MK459915

WPH188 Prophantis cf. androstigmata (Hampson, 1918) Trichaeini (NCBI) (NCBI) JX017853 JX017853 JX017930 —

ZMBN017 Prophantis xanthomeralis (Hampson, 1918) Trichaeini Angola MTD MK459731 MK459916

MTD650 Pycnarmon pantherata (Butler, 1878) Agroterini China MTD MK459732 MK459917

MTD774 Rhectosemia multifarialis Lederer, 1863 Lineodini Venezuela MHNG JF852398 MK459733 MK459918

MTD874 Rhimphalea cf. astrigalis Hampson, 1899 Margaroniini Philippines MTD MK459734 MK459919

ZMBN011 Salbia haemorrhoidalis (Guenée, 1854) Spilomelini Bolivia MTD MK506086 MK459735 MK459920

MTD1358 Samea cf. multiplicalis (Guenée, 1854) Nomophilini Bolivia MTD MK506079 MK459736 MK459921

MTD1235 Samea ecclesialis Guenée, 1854 Nomophilini USA FMNH MK459737 MK459922

MTD872 Siga liris (Cramer, 1775) Spilomelini French Guiana MHNG MK459738 MK459923

MTD1248 Spilomela perspicata (Fabricius, 1787) Spilomelini Peru ZSM MK506084 MK459739 MK459924

MTD783 Spoladea recurvalis (Fabricius, 1775) Hymeniini Morocco MTD MK459740 MK459925

MTD1320 Syllepis marialis Poey, 1832 Hydririni Bolivia MTD LR135741 LR135741 MK459926

MTD1017 Syngamia ﬂorella (Stoll in Cramer & Stoll, 1781) Spilomelini Peru ZSM MK506076 MK459741 MK459927

MTD1315 Terastia meticulosalis Guenée, 1854 Margaroniini Peru ZSM MK506111 MK459742 MK459928

MTD1247 Trichaea pilicornis Herrich-Schäer, 1866 Trichaeini Peru ZSM MK506089 MK459743 MK459929

MTD870 Udea ferrugalis (Hübner, 1796) Udeini Morocco MTD JF852252 MK459744 MK459930

MTD956 Udea washingtonalis (Grote, 1882) Udeini Canada MTD MK459745 MK459931

MTD276,

MTD357 Udeoides muscosalis (Hampson, 1913) Udeini Kenya MTD JF497033 JF497033 MK459932

MTD1467 Zebronia phenice (Stoll in Cramer & Stoll, 1782) Margaroniini Angola MTD MK459746 MK459933

MTD1338 Achyra cf. rantalis (Guenée, 1854) Pyraustini Bolivia MTD MK506109 MK459747 MK459934

MM01851 Anania hortulata (Linnaeus, 1758) Pyraustini (NCBI) (NCBI) GU828675 GU828472 GU829003 GU829287

MTD553 Anania verbascalis (Denis & Schiermüller, 1775) Pyraustini Romania MTD MK459748 MK459935

MTD1484 Cryptosara caritalis (Walker, 1859) Portentomorphini Angola MTD MK459749 MK459936

MTD1558 Euclasta gigantalis Viette, 1957 Euclastini Kenya NHMO MK459750 MK459937

MTD1466 Euclasta splendidalis (Herrich-Schäer, 1848) Euclastini Bulgaria S. Beshkov MK459751 MK459938

MTD1327

(WPH215) Hyalobathra crenulata Sutrisno & Horak, 2003 Portentomorphini Australia UHIM JX017826 JX017826 JX017943 MK459939

MTD1350 Hyalorista cf. taeniolalis (Guenée, 1854) Pyraustini Bolivia MTD MK506108 MK459752 MK459940

MTD605 Loxostege aeruginalis (Hübner, 1796) Pyraustini Macedonia MTD MK459753 MK459941

MTD1343 Oenobotys sp. Pyraustini Bolivia MTD MK506104 MK459754 MK459942

MTD388 Ostrinia nubilalis (Hübner, 1796) Pyraustini Germany MTD MK459755 MK459943

ZMBN096 Pagyda salvalis Walker, 1859 Pyraustini Malaysia ZMBN MK459756 MK459944

MTD906 Paracorsia repandalis (Denis & Schiermüller, 1775) Pyraustini Kyrgyzstan N. Pöll MK459757 MK459945

MTD1322 Portentomorpha xanthialis (Guenée, 1854) Portentomorphini Bolivia MTD MK506077 MK459758 MK459946

MTD477 Psammotis pulveralis (Hübner, 1796) Pyraustini Germany MTD MK459759 MK459947

MTD1344 Pseudopyrausta cf. minima (Hedemann, 1894) Pyraustini Bolivia MTD MK506107 MK459760 MK459948

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

146

Table 1 continued.

DNA sample Genus Species Tribus Origin Collection COI (1440bp) EF-1a (1071bp)

HybLCO/HybNancy HybJerry/HybPat HybOscar-6143/Bosie-6144 HybEF51.9/EFrcM4

MTD953 Pyrausta purpuralis (Linnaeus, 1758) Pyraustini Germany MTD MK459761 MK459949

MTD560 Sitochroa verticalis (Linnaeus, 1758) Pyraustini Romania MTD MK459762 MK459950

MTD1326

(WPH209) Tetridia caletoralis (Walker, 1859) Tetridiini Australia UHIM JX017861 JX017861 JX017938 MK459951

WPH054 Uresiphita gilvata (Fabricius, 1794) unplaced Hawaii UHIM JX017825 JX017825 JX017919 —

DNA sample Genus Species Tribus Origin Collection CAD (825bp) GAPDH (654bp) IDH (657bp) RpS5 (576bp)

HybCAD743f/

HybCAD1028r

HybFrigga/

HybBurre

HybIDHdeg27F/

HybIDHdegR

HybRpS5f/

HybRpS5r

MTD393 Synaphe punctalis (Fabricius, 1775) (PYRALINAE) Germany MTD MK459763 MK460136 MK459952 MK460052

MM00051 Pyralis farinalis (Linnaeus, 1758) (PYRALINAE) (NCBI) (NCBI) GU828092 GU829747 GU829979 GU830604

MM09194 Schoenobius gigantellus (Denis & Schiermüller, 1775) (SCHOENOBIINAE) (NCBI) (NCBI) GU828306 GU829903 GU830222 GU830790

MM11223 Clepsicosma iridia Meyrick, 1888 (ACENTROPINAE) (NCBI) (NCBI) GU828315 GU829906 GU830230 GU830800

MM03362 Crambus uliginosellus Zeller, 1850 (CRAMBINAE) (NCBI) (NCBI) GU828182 GU829811 GU830078 GU830681

MM04967 Eudonia truncicolella (Stainton, 1849) (SCOPARIINAE) (NCBI) (NCBI) GU828197 GU829823 GU830095 GU830697

MTD370 Midila guianensis Munroe, 1970 (MIDILINAE) French Guiana MTD MK459764 MK460137 MK459953 MK460053

MTD1307 Sufetula diminutalis (Walker, 1866) (LATHROTELINAE) Germany MTD MK459765 MK460138 MK459954 MK460054

ZMBN094 Aetholix cf. ﬂavibasalis (Guenée, 1854) Agroterini Malaysia ZMBN MK459766 MK460139 MK459955 MK460055

MTD1016 Agathodes designalis Guenée, 1854 Margaroniini Peru ZSM MK459767 — MK459956 MK460056

MTD1328

(WPH221) Agrioglypta excelsalis (Walker, 1866) Margaroniini Australia UHIM JX017793 — MK459957 JX018024

MTD488A Agrotera nemoralis (Scopoli, 1863) Agroterini Germany MTD MK459768 MK460140 MK459958 MK460057

MTD1354 Anageshna cf. primordialis (Dyar, 1906) Steniini Bolivia MTD MK459769 MK460141 MK459959 MK460058

MTD798 Antigastra catalaunalis (Duponchel, 1833) Margaroniini Morocco MTD MK459770 — MK459960 MK460059

MTD1331 Apilocrocis novateutonialis Munroe, 1968 Wurthiini Peru ZSM MK459771 — MK459961 —

MTD668 Aristebulea principis Munroe & Mutuura, 1968 Wurthiini China MTD MK459772 MK460142 MK459962 —

MTD797 Arnia nervosalis (Guenée, 1854) Nomophilini Morocco MTD MK459773 — — MK460060

MTD776 Arthromastix lauralis (Walker, 1859) Asciodini Venezuela MHNG — MK460143 MK459963 MK460061

MTD1061 Arthromastix pactolalis (Guenée, 1854) Asciodini French Guiana R. Rougerie MK459774 MK460144 MK459964 MK460062

MTD1325 Asciodes cf. gordialis Guenée, 1854 Asciodini Bolivia MTD MK459775 MK460145 MK459965 MK460063

MTD1019 Asturodes ﬁmbriauralis (Guenée, 1854) Margaroniini Peru ZSM MK459776 — MK459966 MK460064

MTD1347 Ategumia ebulealis (Guenée, 1854) Nomophilini Bolivia MTD MK459777 — MK459967 MK460065

MTD1329 Azochis cf. ruﬁdiscalis Hampson, 1904 Margaroniini Peru ZSM MK459778 MK460146 MK459968 MK460066

MTD882 Bocchoris cf. inspersalis (Zeller, 1852) Nomophilini Sierra Leone T. Karisch — MK460147 — MK460067

MTD1281 Botyodes diniasalis (Walker, 1859) Margaroniini China MTD — — MK459969 MK460068

MTD1319 Cadarena pudoraria (Hübner, 1825) Margaroniini Cameroon A. Zwick MK459779 MK460148 MK459970 MK460069

MTD826 Cnaphalocrocis cf. medinalis (Guenée, 1854) Spilomelini Philippines MTD MK459780 — MK459971 MK460070

MTD1041 Conchylodes zebra (Sepp, 1850) Udeini French Guiana MTD MK459781 MK460149 MK459972 —

ITBC058 Conogethes pandamalis (Walker, 1859) Margaroniini Malaysia ZMBN MK459782 MK460150 MK459973 MK460071

147

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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Table 1 continued.

DNA sample Genus Species Tribus Origin Collection CAD (825bp) GAPDH (654bp) IDH (657bp) RpS5 (576bp)

HybCAD743f/

HybCAD1028r

HybFrigga/

HybBurre

HybIDHdeg27F/

HybIDHdegR

HybRpS5f/

HybRpS5r

MTD649 Cydalima perspectalis (Walker, 1859) Margaroniini China MTD — MK460151 MK459974 —

MTD1047 Desmia cf. tages (Cramer, 1777) Nomophilini French Guiana MTD MK459783 MK460152 MK459975 MK460072

MTD1323 Diaphania hyalinata (Linnaeus, 1767) Margaroniini Bolivia MTD MK459784 MK460153 MK459976 MK460073

MTD557 Diasemia reticularis (Linnaeus, 1761) Nomophilini Romania MTD MK459785 MK460154 MK459977 MK460074

MTD1357 Diasemiopsis leodocusalis (Walker, 1859) Nomophilini Bolivia MTD MK459786 MK460155 MK459978 MK460075

ZMBN097 Dichocrocis cf. zebralis (Moore, 1867) Margaroniini Malaysia ZMBN MK459787 MK460156 MK459979 MK460076

MTD868 Dolicharthria punctalis (Denis & Schiermüller, 1775) Steniini Spain MTD MK459788 — MK459980 MK460077

MTD786 Duponchelia fovealis Zeller, 1847 Steniini Morocco MTD MK459789 MK460157 MK459981 MK460078

MTD1316 Eporidia dariusalis Walker, 1859 Spilomelini Cameroon A. Zwick MK459790 — MK459982 MK460079

MTD1321 Eurrhyparodes cf. lygdamis Druce, 1902 Herpetogrammatini Bolivia MTD MK459791 MK460158 MK459983 MK460080

MTD1566 Filodes sp. Margaroniini Angola MTD MK459792 — MK459984 MK460081

MTD1318 Ghesquierellana cf. hirtusalis (Walker, 1859) Margaroniini Cameroon A. Zwick MK459793 MK460159 MK459985 MK460082

MTD1285 Glyphodes sibillalis Walker, 1859 Margaroniini Peru ZSM MK459794 MK460160 MK459986 MK460083

MTD820 Glyphodes cf. stolalis Guenée, 1854 Margaroniini Philippines MTD MK459795 MK460161 MK459987 MK460084

MTD1283 Gonocausta sp. Hydririni Bolivia MTD MK459796 MK460162 MK459988 MK460085

MTD808 Haritalodes derogata (Fabricius, 1775) Agroterini Philippines MTD MK459797 MK460163 MK459989 —

MTD994 Herpetogramma phaeopteralis (Guenée, 1854) Herpetogrammatini Peru ZSM MK459798 — MK459990 MK460086

MTD1337 Hileithia cf. obliqualis (Schaus, 1912) Herpetogrammatini Peru ZSM MK459799 MK460164 MK459991 MK460087

MTD1282 Hodebertia testalis (Fabricius, 1794) Margaroniini Yemen MTD MK459800 MK460165 MK459992 MK460088

MTD1565 Hydriris ornatalis (Duponchel, 1832) Hydririni Angola MTD MK459801 MK460166 — —

MTD1004 Hymenia perspectalis (Hübner, 1796) Hymeniini Peru ZSM MK459802 MK460167 MK459993 MK460089

MTD1043 Lamprosema cf. dorisalis (Walker, 1859) Hydririni French Guiana MTD MK459803 MK460168 MK459994 MK460090

MTD1562 Leucinodes africensis Mally et al., 2015 Lineodini Angola MTD MK459804 — MK459995 —

MTD1349 Leucochroma corope (Stoll in Cramer & Stoll, 1781) Margaroniini Bolivia MTD MK459805 MK460169 MK459996 MK460091

MTD1251 Lineodes vulniﬁca Dyar, 1913 Lineodini Bolivia MTD MK459806 MK460170 MK459997 MK460092

MTD1284 Liopasia andrealis Dognin, 1910 Margaroniini Bolivia MTD MK459807 MK460171 MK459998 MK460093

WPH197 Marasmia poeyalis (Boisduval, 1833) Spilomelini (NCBI) (NCBI) JX017781 — — JX018009

WPH115 Marasmia trapezalis (Guenée, 1854) Spilomelini (NCBI) (NCBI) JX017777 — — JX018002

MTD1341 Maruca vitrata (Fabricius, 1787) Margaroniini Bolivia MTD MK459808 — MK459999 MK460094

MTD364 Mecyna lutealis (Duponchel, 1833) Nomophilini Italy TLMF — MK460172 MK460000 MK460095

MTD1340 Megastes cf. pusialis Snellen, 1875 Margaroniini Bolivia MTD MK459809 MK460173 MK460001 MK460096

MTD787 Metasia suppandalis (Hübner, 1823) Steniini Morocco MTD MK459810 MK460174 MK460002 MK460097

ZMBN104 Nacoleia insolitalis (Walker, 1862) Margaroniini Malaysia ZMBN MK459811 — MK460003 —

ZMBN103 Neoanalthes cf. pseudocontortalis Yamanaka & Kirpichnikova, 1993 Agroterini Malaysia ZMBN — MK460175 MK460004 MK460098

MTD1046 Neoleucinodes dissolvens (Dyar, 1914) Lineodini French Guiana MTD MK459812 MK460176 MK460005 —

MTD152 Niphopyralis chionesis Hampson, 1919 Wurthiini Australia ANIC MK459813 — MK460006 MK460099

MTD782 Nomophila noctuella (Denis & Schiermüller, 1775) Nomophilini Morocco MTD MK459814 MK460177 MK460007 MK460100

MTD1483 Obtusipalpis pardalis Hampson, 1896 Margaroniini Angola MTD MK459815 MK460178 MK460008 MK460101

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

148

Table 1 continued.

DNA sample Genus Species Tribus Origin Collection CAD (825bp) GAPDH (654bp) IDH (657bp) RpS5 (576bp)

HybCAD743f/

HybCAD1028r

HybFrigga/

HybBurre

HybIDHdeg27F/

HybIDHdegR

HybRpS5f/

HybRpS5r

SDA008A Omiodes continuatalis (Wallengren, 1860) Margaroniini Hawaii UHIM — MK460179 MK460009 —

WPH252B Omiodes humeralis Guenée, 1854 Margaroniini Costa Rica UHIM JX017808 MK460180 MK460010 JX018040

MTD784 Palpita vitrealis (Rossi, 1794) Margaroniini Morocco MTD MK459816 MK460181 MK460011 MK460102

MM00325 Patania ruralis (Scopoli, 1763) Agroterini (NCBI) (NCBI) GU828133 GU829772 GU830021 GU830638

MTD1018 Patania cf. silicalis (Guenée, 1854) Agroterini Peru ZSM MK459817 MK460182 MK460012 MK460103

MTD1324 Phostria cf. tedea (Stoll in Cramer & Stoll, 1780) Agroterini Bolivia MTD MK459818 MK460183 MK460013 MK460104

MTD1033 Prenesta cf. iphiclalis (Walker, 1859) Margaroniini French Guiana MTD MK459819 MK460184 MK460014 MK460105

MTD1342 Prenesta cf. rubrocinctalis (Guenée, 1854) Margaroniini Bolivia MTD MK459820 — MK460015 MK460106

MTD1015 Prenesta scyllalis (Walker, 1859) Margaroniini Peru ZSM MK459821 MK460185 MK460016 MK460107

WPH188 Prophantis cf. androstigmata (Hampson, 1918) Trichaeini (NCBI) (NCBI) — — — JX018006

ZMBN017 Prophantis xanthomeralis (Hampson, 1918) Trichaeini Angola MTD — — MK460017 MK460108

MTD650 Pycnarmon pantherata (Butler, 1878) Agroterini China MTD MK459822 MK460186 MK460018 MK460109

MTD774 Rhectosemia multifarialis Lederer, 1863 Lineodini Venezuela MHNG MK459823 MK460187 MK460019 —

MTD874 Rhimphalea cf. astrigalis Hampson, 1899 Margaroniini Philippines MTD — MK460188 MK460020 MK460110

ZMBN011 Salbia haemorrhoidalis (Guenée, 1854) Spilomelini Bolivia MTD — MK460189 MK460021 MK460111

MTD1358 Samea cf. multiplicalis (Guenée, 1854) Nomophilini Bolivia MTD — MK460190 MK460022 MK460112

MTD1235 Samea ecclesialis Guenée, 1854 Nomophilini USA FMNH MK459824 MK460191 MK460023 MK460113

MTD872 Siga liris (Cramer, 1775) Spilomelini French Guiana MHNG MK459825 — MK460024 MK460114

MTD1248 Spilomela perspicata (Fabricius, 1787) Spilomelini Peru ZSM MK459826 — MK460025 MK460115

MTD783 Spoladea recurvalis (Fabricius, 1775) Hymeniini Morocco MTD MK459827 MK460192 MK460026 MK460116

MTD1320 Syllepis marialis Poey, 1832 Hydririni Bolivia MTD LR134539 LR134626 LR134717 LR134887

MTD1017 Syngamia ﬂorella (Stoll in Cramer & Stoll, 1781) Spilomelini Peru ZSM MK459828 MK460193 MK460027 MK460117

MTD1315 Terastia meticulosalis Guenée, 1854 Margaroniini Peru ZSM — — MK460028 MK460118

MTD1247 Trichaea pilicornis Herrich-Schäer, 1866 Trichaeini Peru ZSM MK459829 MK460194 MK460029 MK460119

MTD870 Udea ferrugalis (Hübner, 1796) Udeini Morocco MTD MK459830 MK460195 MK460030 —

MTD956 Udea washingtonalis (Grote, 1882) Udeini Canada MTD — MK460196 MK460031 —

MTD276,

MTD357 Udeoides muscosalis (Hampson, 1913) Udeini Kenya MTD MK459831 MK460197 MK460032 —

MTD1467 Zebronia phenice (Stoll in Cramer & Stoll, 1782) Margaroniini Angola MTD MK459832 MK460198 MK460033 MK460120

MTD1338 Achyra cf. rantalis (Guenée, 1854) Pyraustini Bolivia MTD MK459833 MK460199 MK460034 MK460121

MM01851 Anania hortulata (Linnaeus, 1758) Pyraustini NCBI NCBI GU828170 GU829798 GU830062 GU830669

MTD553 Anania verbascalis (Denis & Schiermüller, 1775) Pyraustini Romania MTD MK459834 MK460200 MK460035 MK460122

MTD1484 Cryptosara caritalis (Walker, 1859) Portentomorphini Angola MTD MK459835 — MK460036 MK460123

MTD1558 Euclasta gigantalis Viette, 1957 Euclastini Kenya NHMO MK459836 — MK460037 MK460124

MTD1466 Euclasta splendidalis (Herrich-Schäer, 1848) Euclastini Bulgaria S. Beshkov MK459837 — MK460038 MK460125

MTD1327

(WPH215) Hyalobathra crenulata Sutrisno & Horak, 2003 Portentomorphini Australia UHIM JX017788 — MK460039 JX018019

149

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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Table 1 continued.

DNA sample Genus Species Tribus Origin Collection CAD (825bp) GAPDH (654bp) IDH (657bp) RpS5 (576bp)

HybCAD743f/

HybCAD1028r

HybFrigga/

HybBurre

HybIDHdeg27F/

HybIDHdegR

HybRpS5f/

HybRpS5r

MTD1350 Hyalorista cf. taeniolalis (Guenée, 1854) Pyraustini Bolivia MTD MK459838 — MK460040 MK460126

MTD605 Loxostege aeruginalis (Hübner, 1796) Pyraustini Macedonia MTD MK459839 MK460201 MK460041 MK460127

MTD1343 Oenobotys sp. Pyraustini Bolivia MTD MK459840 — MK460042 MK460128

MTD388 Ostrinia nubilalis (Hübner, 1796) Pyraustini Germany MTD MK459841 — MK460043 MK460129

ZMBN096 Pagyda salvalis Walker, 1859 Pyraustini Malaysia ZMBN MK459842 MK460202 MK460044 MK460130

MTD906 Paracorsia repandalis (Denis & Schiermüller, 1775) Pyraustini Kyrgyzstan N. Pöll MK459843 MK460203 MK460045 MK460131

MTD1322 Portentomorpha xanthialis (Guenée, 1854) Portentomorphini Bolivia MTD MK459844 — MK460046 MK460132

MTD477 Psammotis pulveralis (Hübner, 1796) Pyraustini Germany MTD — MK460204 MK460047 —

MTD1344 Pseudopyrausta cf. minima (Hedemann, 1894) Pyraustini Bolivia MTD MK459845 MK460205 MK460048 MK460133

MTD953 Pyrausta purpuralis (Linnaeus, 1758) Pyraustini Germany MTD MK459846 MK460206 MK460049 MK460134

MTD560 Sitochroa verticalis (Linnaeus, 1758) Pyraustini Romania MTD MK459847 MK460207 MK460050 MK460135

MTD1326

(WPH209) Tetridia caletoralis (Walker, 1859) Tetridiini Australia UHIM JX017785 — MK460051 JX018014

WPH054 Uresiphita gilvata (Fabricius, 1794) unplaced Hawaii UHIM JX017770 — — JX017996

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.

2. Material and methods

2.1. Material

A broad range of Spilomelinae taxa was investigated to re-

ect the morphological, ecological, evolutionary, and geo-

graphical diversity of the group. 86 Spilomelinae species of

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: Eudonia truncicolella (Stain-

ton, 1849) (Scopariinae), Crambus uliginosellus Zeller, 1850

(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 Pyralidae Synaphe punctalis (Fabricius, 1775) and

Pyralis farinalis (Linnae us, 1758) (both Pyralinae). See Table

1 for the list of taxa that were studied both genetically and

morphologially. 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 identied to ge-

nus- or species level based on morphological investigations

including genitalia dissection, as well as comparing the 5’ half

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

lack by coding morphological features based on information

from published literature (illustrations, descriptions). These

cases concerned the following taxa and consulted literature:

Midila guianensis (Munroe 1970), Diasemiopsis leodocusalis

(walker, 1859) (Munroe 1957), Neoleucinodes dissolvens

(dYar, 1914) (capps 1948), Euclasta gigantalis Viette, 1957

(po pescu-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

the available genitalia, and where possible, COI barcode data.

Those cases with replacement specimens are (male / female): An-

ageshna primordalis / A. cf. primordalis; Asciodes cf. gor dia-

lis / A. quietalis; Azochis cf. rudiscalis / A. rudisca lis; Me -

gastes cf. pusialis / M. pusialis; Trichaea pilicornis / T. pro-

chyta; Hyalorista cf. taeniolalis / H. taeniolalis.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

150

We state author and year of description of all genera

and species at their rst mention in the text. Taxa identi-

ed as ‘cf.’ have the author and year of description of the

closest species known to us. A list of investigated genita-

lia slides is given in Supplement Table S1.

2.2. Methods

2.2.1. Molecular methods

DNA extraction was done using the DNeasy Blood &

Tissue kit (Qiagen) or the NucleoSpin Tissue kit (Mach-

erey-Nagel) according to the manufacturers’ protocols.

The six genes COI, CAD, EF-1a, GAPDH, IDH and

RpS5 were amplied with the following primer pairs:

COI in one large fragment with HybLCO (forward)

and HybPat (reverse) or as two shorter fragments with

HybLCO (forward) and HybNancy (reverse) as well as

HybJerry (forward) and HybPat (reverse); CAD with

HybCAD743f (forward) and HybCAD1028r (reverse);

EF-1a (Elongation Factor 1-alpha) in one large frag-

ment with HybOscar-6143 (forward) and HybEFrcM4

(reverse) or as two shorter, overlapping fragments with

HybOscar-6143 (forward) and HybBosie-6144 (reverse)

as well as HybEF51.9 (forward) and HybEFrcM4 (re-

verse); GAPDH with HybFrigga (forward) and Hyb-

Burre (reverse); IDH with HybIDHdeg27F (forward)

and HybIDHdegR (reverse); RpS5 with HybRpS5f (for-

ward) and HybRpS5r (reverse) (waHlberg & wHeat

2008; Haines & rubinoff 2012). Each primer contains

a universal T7 (forward) or T3 (reverse) primer tail at

their 5’ end, which was used for sequencing (waHlberg

& wHeat 2008).

All gene fragments were amplied in 25 µl reactions.

The amplication protocol at the SNSD DNA lab was:

200 nM of each primer, 200 µM dNTP mix, 2.5 µl Taq

buffer, 1 mM MgCl2, 1 u BIO-X-ACT Short DNA Poly-

merase (Bioform), 2 µl DNA of concentration as extract-

ed, and distilled water added up to 25 µl in total per reac-

tion. At the UiB DNA lab the PCR protocol was: 400nM

of each primer, 800 µM dNTP mix, 2.5 µl Taq buffer

(incl. MgCl2), 0.75 u TaKaRa Ex Taq DNA Polymerase,

2 µl DNA of extracted concentration, and distilled water

added up to 25 µl in total per reaction.

The PCR programme for mitochondrial COI was: ini-

tial phase at 95°C for 5 min, 38 – 40 cycles at 95°C for

30 s 50°C for 30 s and 72°C for 60 s, nal phase at 72°C

for 10 min and cooling at 8°C. For the nuclear genes

CAD, EF-1a, GAPDH, IDH and RpS5 we ran a touch-

down PCR: 24 cycles at 95°C for 30 s 55°C with – 0.4°C

/ cycle for 30 s and 72°C for 60 s + 2 s / cycle, then 12

cycles at 95°C for 30 s 45°C for 30 s and 72°C for 120 s

+ 3 s / cycle, nal phase at 72°C for 10 min and cooling

at 8°C.

PCR results were examined via gel electrophoresis

on a 1% agarose gel and GelRed as dying agent. Suc-

cessful PCR samples were cleaned with ExoSAP and

subsequently amplied in Sanger-sequencing PCR reac-

tions. Sequencing was done in both directions with the

T7 and T3 primers, using the BigDye kit with this setup:

0.5 – 3.0 µl of PCR sample (depending on the sample’s

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

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-

tercycler ep gradient s (Eppendorf) or a PCR System 9700

(GeneAmp), at UiB a Bio-Rad 1000 thermal cycler was

used for PCR and sequencing PCR, and a MJ Research

PTC-200 thermal cycler for PCR clean-up. All sequences

were proofread by eye and aligned manually using PhyDE

0.9971 (Müller et al. 2008). All new sequence data have

been submitted to an open access nucleotide sequence

database (GenBank; https://www.ncbi.nlm.nih.gov/gen-

bank); accession numbers are compiled in Table 1.

2.2.2. Morphological methods

Genitalia were dissected according to robinson (1976),

with modications: The abdomen was cut open along one

pleural membrane, cleaned, and embedded in medium

under a cover slip to allow clear investigation of the tym-

panal organs. Female genitalia were stained with Chlo-

razol Black. Male genitalia were either left unstained or

were stained with Chlorazol Black or Eosin Y.

Morphological structures were investigated using

Leica M125 and M205C stereomicroscopes. Imagines

were photographed with a Canon EOS 60D in combi-

nation with a Canon EF 100mm 1:2,8 Macrolens and

Canon EOS Utility Version 2.10.2.0. A Leica CTR6000

Microscope in combination with a Leica DFC420 camera

and Leica Application Suite programme (Version 3.8.0)

was used to photograph the genitalia.

Observed morphological features were coded accord-

ing to the morphology character circumscriptions and

compiled in a morphomatrix (Table 2) for all investigated

taxa. Clepsicosma iridia (Acentropinae) was not studied

morphologically, and is therefore omitted in Table 2.

Morphological abbreviations in Figs. 3 – 15: an. – antrum; a.t. –

anal tube; ap.a. – apophysis anterioris; ap.p. – apophysis posterio-

ris; apx. – appendix bursae; coe. – coecum of phallus; col. – col-

liculum; cos. – costa; cos.b. – costa base; cos.d. – distal costa; cos.

ex. – dorsad process of basal costa; cos.p. – rod-shaped ventrad

process of basal costa; crn. – cornutus or cornuti; c.b. – corpus

bursae; d.b. – ductus bursae; d.s. – ductus seminalis; div. – diver-

ticulum; fo.ty. – fornix tympani; b. – bula; a. – basal antennal

agellomeres; f r. – frons; fre. – frenulum; gna. – gnathos; h.p. –

hairpencil scerite(s); h.p.s. – hairpencil sclerites of the saccus;

hau. – haustellum; jx. – juxta; la.p. – labial palps; lam. – lamella

antevaginalis of ostium bursae; lob. – lobulus of lateral tympanal

case; mx.p. – maxillary palps; o.b. – ostium bursae; oc. – ocellus;

p.a. – papilla anales; p.ph. – posterior phallus apodeme; ped. –

pedicellus; pl.m. – pleural membranes; pl.tu. – pleural scale tufts

of male abdominal segment 7; pl.sc. – pleural sclerites of male

abdominal segment 8; ret. – retinaculum; s2 – s8 – 2nd – 8th ab-

dominal sternite; sac. – sacculus; sac.d. – distal sacculus; sac. ex. –

151

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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extension of distal sacculus; sac.pr. – projection from central sac-

culus; sc. – saccus of vinculum; sc.v. – ventral saccus tip; sca.f. –

raised scales on mesal side of agellomeres; se.s. – sensillar setae

of agellomeres; sig. – signum; t1 – t8 – 1st – 8th abdominal tergite;

teg. – tegumen; teg.h. – hair-pencils on the dorsolateral tegumen;

tran. – transtillum arm; tr.in. – transtillum inferior sensu Marion

1954; ty.fr. – tympanic frame; unc. – uncus; u.ch. – uncus chaetae;

v.s. – venula secunda; v.va. – ventral valva edge; ves. – vesica;

vin. – vinculum; vin.d. – dorsal joint of vinculum with valva.

2.2.3. Phylogenetic analysis

For the phylogenetic analyses, the nucleotide sequences

of the genetic data were used. Initial Maximum Likeli-

hood analyses showed that analysis of the amino acid

sequences translated from the nucleotide data resulted in

poorly resolved topologies with branch supports mostly

>> 50 percent; amino acid sequences were therefore dis-

regarded as phylogenetic data source.

The sequence data were investigated for potential

substitution saturation in third codon positions (Xia et al.

2003; Xia & leMeY 2009) using DAMBE5 (Xia 2013).

RogueNaRok (aberer et al. 2013) was used to screen the

molecular data for rogue taxa, which were subsequently

excluded from the analysis.

We analysed the concatenated molecular and mor-

phological data with two different partitioning schemes:

GENES and TIGER. In the GENES scheme, we placed

each gene and the morphological data into a separate

partition, resulting in 7 partitions. In the TIGER scheme,

following rota & waHlberg (2012), we partitioned the

molecular dataset in terms of evolutionary site rates using

the programme TIGER (cuMMins & McinerneY 2011).

We chose initial partitioning into 10 bins and pooled the

bins with < 100 sites with bin 1, resulting in 5 molecular

partitions, with bin 1 (incl. bins 2 – 6, each with < 100

sites) = 1798 sites, bin 7 = 182 sites, bin 8 = 684 sites, bin

9 = 1291 sites, and bin 10 = 1213 sites; morphology was

treated as a separate 6th partition.

We used jModeltest v2.1.4 (guindon & gascuel 2003;

darriba et al. 2012) to infer the models that best reect

the sequence evolution of the genetic data. The resulting

models for the GENES partitioning are: TIM3+G+I mod-

el for the COI partition, SYM+G+I model for the EF-1a

partition, TVM+G+I model for the GAPDH partition, and

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-

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

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

was analysed with MrBayes version 3.2.6 (ronquist et

al. 2012) on the CIPRES online platform (Miller et al.

2010) using Extreme Science and Engineering Discovery

Environment (XSEDE). Two parallel runs were set up

for 30 Mio. generations, with sampling of every 1,000th

generation. The parameters for gamma shape, proportion

of invariable sites, character state frequencies and GTR

substitution rates were unlinked for the partitions, and the

overall rate was allowed to vary across partitions. The ini-

tial 25% of the trees were discarded as burn-in. Effective

sampling sizes (ESS) and the degree of convergence of

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).

In addition, a Maximum Likelihood (ML) analysis of

the gene-partitioned molecular dataset was done using

RAxML-HPC2 (8.2.10) (staMatakis 2014) on XSEDE

through the CIPRES V 3.3 online platform (Miller et al.

2010).

WinClada 1.00.08 (niXon 2002) was used to derive

ancestral morphological characters from the topology

that was observed in the majority of analyses. Unambig-

uous synapomorphies, and in addition those derived from

slow optimization (slow optimization or DELTRAN,

swofford & Maddison 1987) are plotted on the topology

and summarized. Apomorphies are included in the diag-

noses of clades (see Phylogenetic classication section)

except if they are very homoplastic in that clade. Char-

acters were mapped on the consensus of the parsimony

cladograms as well as the Baysian trees for the sake of

methodological consistency (assis 2015).

A parsimony analysis was conducted with TNT 1.5

(goloboff & catalano 2016). All states were non-addi-

tive and equally weighted, and gaps were treated as miss-

ing data. A traditional search plus the parsimony ratchet

(niXon 1999) and branch-swapping was done (com-

mands: mxram 100; cc-.; collapse [; rs 1; hold 10000; rat:

iter 50; mu: hold 20 replic 100 rat; bb;). Ratchet com-

mands were the default values: stop when 14 substitu-

tions made, 4% upweight and downweight probability,

50 total iterations, alternating equal weights. One hun-

dred replications were done, saving 20 trees per repli-

cation. In addition to equal weights, implied weighting

(goloboff 1993) was explored under a range of k-pa-

rameter values in TNT with the same search parameters.

To try to resolve incongruence among cladograms, we

ran the IterPCR script provided by pol & escapa (2009).

This script suggests characters to recode, which is not

done by the application embedded in TNT 1.5.

3. Results

3.1. Molecular data

We present new genetic data for 100 taxa. In addition,

we complemented the genetic data for four taxa from

the study of Haines & rubinoff (2012), for which we

obtained the original DNA extracts from Will Haines

(University of Hawaii): vouchers WPH209, WPH215,

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

152

WPH221, and WPH252. COI sequencing was successful

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 =

100 samples), for the 1st part of EF-1a 94% (n = 100),

for the scond part of EF-1a 100% (n = 104), for GAPDH

70% (n = 104), for IDH 97% (n = 104), and for RpS5 85%

(n = 100). The amplication of RpS5 with the primers

of waHlberg & wHeat (2008) failed for all taxa in the

tribes Udeini and Lineodini (see Taxonomy for tribes)

except for 37% of the sequence length of Lineodes vul-

nica, probably due to a lack of match between primer

and attachment sequence.

The molecular alignment has a length of 5,223 base-

pairs (bp), with 1,440 bp accounting for COI, 825 bp for

CAD, 1,071 bp for EF-1a, 654 bp for GAPDH, 657 bp

for IDH, and 576 bp for RpS5.

The CAD sequence of Anania verbascalis (GenBank

accession no. MK459834) lacks three codons (9 bp, i.e.

three amino acids in the respective protein product) com-

pared to all other CAD sequences incorporated in our

dataset. These three codons are present in the congeneric

species, A. hortulata, and they code for the amino acids

Isoleucine-Alanine-Valine. This three-codon deletion is

situated in a variable region of the CAD gene, where es-

pecially the second codon is coding for a variety of dif-

ferent amino acids among the investigated taxa. A three-

codon deletion at the identical location in the CAD se-

quence was observed in other Pyraustinae taxa believed

to be closely related to Anania (Kai Chen, pers. comm.),

so that this deletion might represent a synapomorphy for

these taxa.

The long terminal branch of Niphopyralis chionesis

Hampson, 1919 in the phylogenetic results of Mutanen

et al. (2010) and regier et al. (2012) is suspicious, and

we therefore re-sequenced this species from available

material (voucher no. MTD152) to evaluate whether the

data from the previous studies might be compromised.

Our resulting sequence coverage was comparable to that

of Mutanen et al. (2010), where the rst half of EF-1a

and the entirety of the GAPDH were unsuccessful in

sequencing, just as in our results. Our sequenced data

largely matches that of the Mutanen et al. (2010) dataset,

with only a few nucleotide differences between the cor-

responding sequences of the two investigated specimens.

This result strongly suggests that the long terminal branch

of N. chionesis in former studies is not due to sequence

contamination. However, since the same DNA lab proto-

col (waHlberg & wHeat 2008) was used in the Mutanen

et al. (2010) study as well as in the present study, the

sequencing of pseudogenes cannot be ruled out, although

no reading frame shifts or stop codons occur in any of

the investigated N. chionesis sequences, suggesting that

they code for functional proteins. None of the nucleotide

sequences of N. chionesis is found to be exceptionally

divergent from those of other investigated taxa, and ob-

served nucleotide substitutions relative to the other taxa

mostly result in synonymous amino acid codons, i.e. they

encode the same amino acid.

3.2. Morphological data

Morphological data was coded from investigation of dried

adult specimens and their genitalia. For a complete list of

genitalia slides of species investigated in the context of

this study (beyond the taxa included in the phylogenetic

dataset), see Electronic Supplement File 1.

The morphological investigation resulted in the recog-

nition of 115 variable characters for all 114 taxa. Of these

characters, 91 are binary, and 24 are multistate. Nineteen

characters code features of the head and thorax including

legs and wings, 23 of the abdomen including the tym-

panal organs but excluding the genitalia, 47 of the male

genitalia, 25 of the female genitalia and one character of

the locality of larval feeding. Character 115 (locality of

larval feeding) was coded from literature data, and the

following literature was used: HinckleY (1964), gentY &

Mariau (1975), Munroe (1976), allYson (1984), coM-

Mon (1990), nuss (2005), speidel (2005), slaMka (2008,

2013), HaYden et al. (2013), leraut (2014) and pereira

et al. (2014), as well as a personal observation of Leuci-

nodes africensis Mally et al., 2015 from Marja van der

Straten (pers. comm.).

The morphomatrix is shown in Table 2. The deni-

tions of the morphological characters and their states are

as follows:

1 Presence of anteriad-directed projection medially on

frons: (0) absent (Fig. 11C); (1) present (Fig. 4B).

2 Presence of haustellum: (0) absent (Fig. 6A; roepke

1916: g. 2); (1) present (Fig. 4B).

3 Presence of transverse rim on anterior or mesal face

of pedicellus [male]: (0) absent (Fig. 8F); (1) present

(Fig. 11C).

4 Presence of a crest or prong of raised scales on mesal

side of agellomeres [male]: (0) absent (Fig. 11C);

(1) present at ca. 1/3 of antenna length, crest forming

a triangular prong proximally (Fig. 11D); (2) present

in proximal part of antenna (Fig. 8F).

5 Length of sensillar setae at basal antennomeres rela-

tive to diameter of basal antennomeres [male]: (0) ≤

50% (♂ in Fig. 10I, Fig. 11C); (1) > 50% (♂ in Fig.

10H, Fig. 11D).

6 Length of cilia at antenna base in female compared

to male: (0) of equal length (Fig. 10I); (1) shorter

(Fig. 10H).

7 Presence of ocelli: (0) absent (Fig. 6B); (1) present

(Fig. 11C).

8 Direction of 3rd labial palpomere: (0) dorsal (Fig.

11C); (1) porrect (Fig. 4B).

9 Intersexual size differences of 3rd labial palpomere:

(0) well developed in both sexes (Maes 1995: pl. 5);

(1) short in the male (Fig. 4B); (2) short in both sexes

(Fig. 6A; roepke 1916: g. 3).

10 Length of maxillary palpi: (0) long enough to hypo-

thetically come in contact with each other (Fig. 11C);

(1) minute to obsolete, cannot hypothetically get in

contact with each other (Fig. 6A).

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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11 Presence of broad scale tuft on distal foreleg tibia

(not to be confused with epiphysis): (0) absent; (1)

present.

12 Presence of tuft of long scales on distal foreleg fe-

mur: (0) absent; (1) present.

13 Presence of a longitudinal groove bearing a hair pen-

cil on male midleg tibia: (0) absent (frolov et al.

2007: g. 1A); (1) present (frolov et al. 2007: g.

1B,C).

14 Presence of tibio-abdominal scale brush [male]: lon-

gitudinal line of thin, spatulate scales on male hind-

leg’s proximal tibia in contact with an area of ventrad

scales on pleural membranes of abdominal segment

2: (0) absent; (1) present (MeY & speidel 2010: gs.

5, 10).

15 Number of apical spurs on hindtibia: (0) 4, a proxi-

mal and a distal pair (Fig. 10F, G); (1) 2, only a dis-

tal pair (as in g. 10F,G, but without proximal spur

pair).

16 Length of metatibial proximal inner spur relative to

tibial segment between this and the distal spur pair:

(0) < 1/2 (distance “d”) (Fig. 10F); (1) ≥ 1/2 (dis-

tance “d”) (Fig. 10G).

17 Presence of eld of enlarged, raised scales on male

central forewing costa: (0) absent (Fig. 7A); (1) pre-

sent (Fig. 9A).

18 Form of retinaculum at costal base of forewing un-

derside in males: (0) simple brush of straight hairs

(Fig. 8I); (1) cuticle protruded as a retinacular hook

(frenulum hook sensu forbes 1926: g. 7; popescu-

gorJ & constantinescu 1977: g. 3b).

19 Number of frenulum bristles in female: (0) one (Fig.

4A); (1) two (popescu-gorJ & constantinescu 1977:

g. 3a).

20 Splitting of praecinctorium: (0) strong (Marion 1954:

g. 2); (1) weak to absent (Marion 1954: g. 1).

21 Presence of lobulus on lateral edge of tympanal case:

(0) absent (Fig. 6C); (1) present (Fig. 8A).

22 Shape of fornix tympani surface: (0) projecting from

the tympanic frame (Minet 1983: “cd.” in g. 30;

Maes 1985: “f.ty.” in pls. 1A, 1D); (1) recessed with-

in the frame (Maes 1985: pl. 1E, “f.ty.” in pl. 2C).

23 Direction of fornix tympani projection: (0) ven-

tral (Minet 1983: “cd.” in g. 30); (1) lateral (see

HaYden 2013: gs. 18, 19, 21, 22).

24 Presence of venulae secundae: (0) absent (Fig. 6C);

(1) present (Fig. 8A).

25 Course of venulae secundae: (0) converging (Fig.

5C); (1) in posterior half parallel or diverging (Fig.

8A).

26 Presence of lateral anteriad lobe on each side of ante-

rior edge of male sternite 3: (0) absent (Fig. 6C); (1)

present (Fig. 7G).

27 Presence of pleural scale tufts on each side of the

male abdomen, one small scale tuft anteriorly on

segment 6 and one large scale tuft anteriorly on seg-

ment 7: (0) absent (Fig. 3H); (1) present (Fig. 8H).

28 Presence of pleural scale tufts on each side of the

male abdomen, one large scale tuft anteriorly on

segment 5, one small scale tuft on anterior ends of

segments 6 and 7: (0) absent (Fig. 12G); (1) present

(Fig. 13C).

29 Presence of large, oval pleural scale tufts on each

side of male abdominal segment 7, with a presum-

ably secretory opening in its anterior centre: (0) ab-

sent (Fig. 3H); (1) present (Fig. 14F).

30 Outline of central anterior edge of male sternite 7:

(0) straight to slightly undulate (Fig. 6F); (1) anteri-

orly projecting protuberance or spine (Fig. 3H); (2)

arch-shaped recession (Fig. 8G).

31 Outline of central posterior edge of male sternite

7: (0) straight (Fig. 8H); (1) with pair of posteriad,

curved spines running dorsally of sternite 8 (Fig.

6F); (2) with pair of posteriad lobes (Fig. 15E); (3) a

wide V-shaped recession (Fig. 8E).

32 Outline of anterior edge of male tergite 8: (0) straight

to convex (Fig. 6F); (1) with triangular and straight-

edged or semicircular indentation (Fig. 3H).

33 Sclerotization of male tergite 8: (0) homogenous

(Fig. 9D); (1) heterogenous, i.e. with distinct sclero-

ti zation pattern (Fig. 3H).

34 Sclerotization pattern on male tergite 8: (0) central

longitudinal strip; (1) longitudinal strip, bifurcating

anteriorly into a Y-shape (Fig. 3H).

35 Presence of a eld of setose scales on the anterior

ends of the male tergite 8’s Y-shaped sclerotisation:

(0) absent (Fig. 8H); (1) present (Fig. 11F).

36 Presence of U-shaped sclerotisation on lateral and

anterior edge of male sternite 8: (0) absent (Fig. 9D);

(1) present (Fig. 11F).

37 Presence of an anterolaterad sclerotized lobe on each

side of anterior edge of male sternite 8, running dor-

sad of sternite 7: (0) absent (Fig. 8H); (1) present

(Fig. 6G).

38 Presence of central hair scale patch(es) on anterior

edge of male sternite 8: (0) absent (Fig. 8H); (1) pre-

sent (Fig. 3H).

39 Presence of median U-shaped recession or deep

notch on posterior edge of male sternite 8: (0) absent

(Fig. 3H); (1) present (Fig. 6G).

40 Presence of a sclerite on each pleural membrane of

male segment 8: (0) absent (Fig. 6F); (1) present

(Figs 3H, 9D).

41 Shape of pleural sclerite on male segment 8: (0) slim

longitudinal strip (Fig. 9D); (1) broad semicircle

(Fig. 3H).

42 Presence of a eld of setae anterior on pleurites of

male segment 8: (0) absent (Fig. 3H); (1) present.

43 Presence of uncus: (0) absent (reduced) (Fig. 3A);

(1) present (Fig. 3D).

44 Shape of uncus: (0) conical, non-capitate (Fig. 3D);

(1) capitate (Fig. 3G).

45 Shape of apical uncus: (0) single head (Fig. 3D);

(1) bi- or trifurcate head (Fig. 9E); (2) two separate

heads (Figs. 9G, 11E).

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Table 2. 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).

Character 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111

0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111

Taxon: 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345

Synaphe punctalis 0100111100 000001010? 01?1000000 000?000000 ?010010011 2000010000 ??000????0 00?000010? 10011001B0 1000?0000? 010110???? 000?0

Pyralis farinalis 0100111000 000001010? 01?0?00000 000?000000 ?010010411 0000200000 ??000????0 00?000010? 1001100100 000100000? 0000?0???? 000?E

Crambus uliginosellus 0100001100 00000100?0 1010?00000 001??00000 ?011010421 ???0?????0 ??000????0 01?1000000 10011011B2 0111000110 0000?11100 000?1

Eudonia truncicolella 0100011100 0000010010 1011000000 000??00000 ?011010001 ???0000000 ??000????0 00?0000000 1000?000?0 1001000010 0000?1100? 00120

Midila guianensis 01????10?0 ??????0??? ?????????? ?????????? ??10010?01 ???0000000 ??000????0 00?0000000 10011?010? ?????????? ?????????? ?????

Schoenobius gigantellus 0000111100 0000010000 1011000000 010??10010 ?010010401 ???0000000 ??000????0 00?0000100 10011001B0 1000?00010 1000?0???? 000?2

Sufetula diminutalis 01000?00?0 00000100?1 0011100000 000??00000 ?00??0???? ??0??00000 ??000????0 00?0000200 10011001D2 100110000? 000110???? 000?4

Aetholix cf. ﬂavibasalis 0100111001 0000010010 0001100000 0111010000 ?011110312 4100001001 0000111100 0000000100 1000?101D2 000110000? 0010?0???? 000??

Agathodes designalis 0100001121 0000010010 1001100000 0111010001 001000??10 2100000011 0000122100 0110100200 1000?10120 0001100011 0000?10??? 0011C

Agrioglypta excelsalis 0100011120 0000010010 0001100000 0111010001 0110010301 2100110111 10010200?0 0110100100 1000?101B1 000110000? 0001011100 000?0

Agrotera nemoralis 0100111000 0000010010 0000?00000 000??00000 ?010010310 1100120100 ??001200?0 0011000100 10110111B0 001100000? 0100?1100? 000?0

Anageshna cf. primordialis 0100111000 0000010000 1000?00000 0111010001 0011011112 0000000011 ??000????0 00?0000200 1000?10121 000010000? 0200?0???? 000?4

Antigastra catalaunalis 0100001120 00?000001? 1000?00000 010??00000 ?01101101? 0000010000 ??00100100 0111100200 1000?10122 0001000011 0010?0???? 000?C

Apilocrocis novateutonialis 0100111121 0000010010 0??0?00000 000??00000 ?01001A002 0000200000 ??00110100 0010000000 100110012? ?????????? ?????????? ?????

Aristebulea principis 0100011100 0000010010 0000?00000 000??10000 ?011011002 0000200000 ??00110100 0110000110 1001000112 001100001? 0011011??? 000??

Arnia nervosalis 0100111120 0000000010 0001100000 000??10001 0010011012 000010011? ??00100100 0101000100 1011010110 0001000010 0000?1000? 000??

Arthromastix lauralis 0101111020 0100010010 0000?00000 0111010000 ?011011012 1100020001 1101010100 0121100200 1000?00100 000111000? 0011010??? 000??

Arthromastix pactolalis 01001?10?0 00000100?0 0001100000 0111110000 ?011111012 1100?00101 1100132100 0121001100 1000?001B0 001110000? 0010?10??? 000??

Asciodes cf. gordialis 0101111021 1100010010 0000?00000 0111110001 0011211012 2100120001 1100122100 0121000110 1000?00120 0011?0000? 001??10??? 000?0

Asturodes ﬁmbriauralis 0100001021 0000010010 0000?00000 0111010001 011101120? 2000000111 0001022100 0111100200 1001010120 000110000? 0010?10??? 000??

Ategumia ebulealis 0100001120 00000100?0 0000?00000 0011010001 000??0???2 1000020101 00001220?0 0111001210 1001010112 000110000? 000111105? 000??

Azochis cf. ruﬁdiscalis 01021?11?1 00?00100?0 0001100000 4011010001 0011011011 2000000001 11010230?0 0000100200 1000?101D0 100110000? 0001011122 000??

Bocchoris inspersalis 0100111120 0000010010 0001000000 0111010000 ?00??0???? 0000110011 01000????0 00?0001200 10010111D? 1001?1001? 001101A00? 000?3

Botyodes diniasalis 0100001120 0000010010 1001100000 0010010001 0011011012 2100020011 11000????0 00?1100100 1000?10121 000110000? 001100???? 000?0

Cadarena pudoraria 0100001120 0000010010 0001100000 0111010000 ?011011222 0000?10111 00001200?0 0110100200 0000?101B0 000110000? 0?00?10??? 0010?

Cnaphalocrocis medinalis 0100011120 0000011010 1000?00000 0111010001 001121101? 0010001101 0001020100 0001010100 1000?00120 0001000010 001101100? 000?0

Conchylodes zebra 0100001021 0000000010 0000?00000 010??10000 ?0100101?? 0000120000 ??00120100 0110010100 1001101120 0001000011 000101102? 0012?

Conogethes pandamalis 01000?10?1 00000100?0 0001100001 0111010001 0011010012 2000000001 0101000100 0111110200 1000?101B? ?????????? ?????????? ?????

Cydalima perspectalis 0100001120 0000000010 1001100001 0?11010001 001111A202 2100100111 0100122100 0111100200 1000?101B1 000110000? 0011111100 000?0

Desmia tages 0111011021 0000010010 0001100000 0111010001 001101A012 2000001111 0101020100 0011000200 1000?101D0 000101000? 0010?0???? 000?0

Diaphania hyalinata 0100011120 00000000?0 1001100000 0111010001 0111011022 4000010001 1101030100 0111101200 1000?11121 000110000? 000101A101 000?0

Diasemia reticularis 0100111100 0000010011 0001100000 010??10000 ?00??0???? 0000100000 ??000????0 00?1000200 1001111110 100110000? 0000?1100? 000?0

Diasemiopsis leodocusalis 0100111110 0000010011 1?00?????? ?????????? ??1101101? ??00000000 ??000????0 00?1000200 1001100111 000110000? 001111105? 000??

Dichocrocis cf. zebralis 0100111021 0000010010 1001100000 0111010001 0010010E22 0000110001 00010220?0 0111100000 1000?101D2 0001?10011 1000?11100 000??

Dolicharthria punctalis 0100111100 0000010011 0000?00000 0011010001 0011011012 0000000001 0001032101 0001?00000 1000?00122 000110000? 0010?0???? 000?5

Duponchelia fovealis 0100011021 0000010011 0001100000 010??10001 001101102? 0000000001 00010230?0 0000000201 1001101120 0001?1000? 0210?0???? 000?H

Eporidia dariusalis 0100001021 0000000010 0001100000 000??00000 ?010211000 0000010000 ??01020100 0110000200 1000?00110 001101000? 0010?0???? 0011?

Eurrhyparodes lygdamis 0100111110 0000010010 1??0?00000 000??10000 ?01001101? 0000000110 ??00120100 0110000100 1000?1111? 0001100011 0011110??? 000??

155

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Character 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111

0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111

Taxon: 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345

Filodes fulvidorsalis 0100001020 1001010010 1001100000 0111010001 001000??02 0000000001 1101020000 0111100201 1000?10112 0001100011 0000?0???? 0010?

Ghesquierellana hirtusalis 010???11?1 ????0???10 00011????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????2 100110000? 0000?0???? 000??

Glyphodes sibillalis 0100011120 0001010010 0001100000 0111010001 0?11010222 0000?00001 1100122100 0111101200 1000?101B1 0001000010 0000?10??? 000?0

Glyphodes cf. stolalis0100111120 0000010010 1001100000 0111010001 0011010222 4?00020001 1100120100 0111100200 1000?10100 0001100011 1000?10??? 000??

Gonocausta cf. zephyralis 0100111111 00000100?0 0000?00000 000??00100 ?010011012 0000000000 ??000????0 00?0000000 1000?000?1 000110000? 0000?1103? 0010?

Haritalodes derogata 0100001020 0000010010 0001100000 010??00000 ?010010100 0000001101 0100122100 1001010200 1000?10121 000110000? 0001011100 000?0

Herpetogramma phaeopteralis 0100111120 0000010010 0000?00000 000??10001 0010010012 0000110001 0000120100 0000000200 10010101D0 000110000? 0010?0???? 000?0

Hileithia obliqualis 0100111121 000001001? 0??1000000 010??10000 ?010011010 0000110001 0000120100 0010000100 1000?101D1 000110000? 0001010??? 000??

Hodebertia testalis 0100001121 0000010010 0001100001 0111010001 0011011021 0100010011 100012G101 0110100100 1000?01120 001110000? 0000?0???? 000??

Hydriris ornatalis 0100011011 000001001? 0??0?00000 000??10000 ?00??????? ??01200000 ??00122000 0110010000 10011001B0 001100000? 0011011143 00101

Hymenia perspectalis 0110011100 0000010010 0001100000 000??10001 0011011012 0000200001 11000????0 00?0100100 1000?11100 000110001? 000111105? 000??

Lamprosema cf. dorisalis 01000?10?0 00000100?1 0??1100001 0111010101 1011011001 1000000001 1110103000 0120000B00 1000?001D? ?????????? ?????????? ?????

Leucinodes africensis 0100011111 0000010000 0000?00000 000??10000 ?011011010 2000010010 ??010100?0 0100000000 1001001121 0001?1000? 0000?0???? 000?3

Leucochroma corope 0100011020 0000010010 0000?00000 0111010001 0011011012 0000100001 1101022100 0110100200 0000?10121 000110000? 000100???? 000??

Lineodes vulniﬁca 0100001121 0000000000 0??1100000 000??00000 ?010011002 0000010000 ??001200?0 0010000000 1000?01120 000110000? 0000?0???? 000?A

Liopasia andrealis 0100011100 00000000?0 0001100000 0010010000 ?011010300 0000200000 ??001020?0 0110000100 1001101111 001110000? 0000?0???? 00113

Marasmia poeyalis 0100011120 0000011010 1001100000 01110??001 0011211010 ??00010001 10000????0 01?1000000 1000?101D0 0001000010 001101100? 000?0

Marasmia trapezalis 0100011100 0000010010 00011????? ?????????? ??1121101? ??00010101 00000????0 00?0000100 1000?00112 0001000010 001101100? 000??

Maruca vitrata 0100011101 0000010010 1000?00000 0111010001 001101A102 0000?01101 11010220?0 0110100001 1000?11122 000110000? 000101A111 0011G

Mecyna lutealis 0100111120 0000010010 0001100000 000??10001 0010011012 0000000001 00001200?0 0011000200 1001010110 0001110010 0000?1101? 000?0

Megastes cf. pusialis 0100111100 0000010010 0001100000 000??00000 ?01001A012 0000010010 ??001220?0 0000000200 1000?000?0 000110000? 0000?10??? 000??

Metasia suppandalis 0100011100 0000000000 0001100000 0011010001 001121101? 0000010111 0000122100 0001000100 1001010122 001110000? 001100???? 000??

Nacoleia insolitalis 0100011020 0000010011 0000?00000 0010010001 0011010022 2100?00001 0000120101 0110101200 1000?10120 000110000? 001101100? 000??

Neoanalthes cf. pseudocontortalis 0100111021 0000010010 ?001100000 000??10001 0010010002 4000001101 0000132101 0001000200 1000?10122 0001100010 0011011100 000??

Neoleucinodes dissolvens 0100111101 000001000? ???0?00000 0111010000 ?011011000 0000000000 ??010320?0 0010000000 1000?001B? 0001?0000? 0010?0???? 000??

Niphopyralis chionesis 0000110020 00001?0000 0000?00000 100??01010 ?0112????? ??00?00000 ??000????0 00?0000010 1000?00120 0011?0000? 000100???? 000??

Nomophila noctuella 0100111101 0000010010 0001100000 010??10001 0011110010 1000120001 0000122100 0010000200 1000?10100 1001000110 001101101? 000?0

Obtusipalpis pardalis 0100111100 00000100?0 0001100000 0011000000 ?01001032? 0000000001 1001030100 0010000200 1000?101D2 0001?1000? 000100???? 000??

Omiodes continuatalis 0100001120 0000010010 1001100000 0111010001 001101A222 0000010101 00010220?0 0111100200 1000?10100 000110000? 0000?11100 100??

Omiodes humeralis 0100111020 00?0010010 1001100000 0111010001 001101AE02 2100020011 00010220?0 0110100200 1000?101B0 000110000? 0010?0???? 000??

Palpita vitrealis 0100001120 0000010010 0001100001 0111010001 0011011012 2000010001 11000????0 01?0100200 1000?101D1 0011?0000? 0011011100 000?C

Patania ruralis 0100011001 0000010010 1000?00000 000??00001 0010010012 0100001111 0000122100 0011010200 1001000010 000110000? 0010?11111 000?0

Patania cf. silicalis 0100001021 0000010010 1000?00000 0111010001 0010010012 4100001101 0100120100 0000000100 1000?00121 000100000? 0000?1101? 000??

Phostria temira 010011102? 0000010010 0001110000 010??10001 0010010002 4100020101 1100122100 0010000200 1000?001D0 0001100011 1000?1100? 000??

Prenesta iphiclalis 0100111020 00000?0?10 00011????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????0 0001100010 1000?0???? 000??

Prenesta cf. rubrocinctalis 01001?11?0 00000100?0 0000?00000 0011010001 001101A2?2 0000000001 0000120100 0111000200 1000?0010? ?????????? ?????????? ?????

Prenesta scyllalis 0101111021 0000010010 0001100000 0111010001 0011011322 2100?0?101 1100102100 0111100200 1000?101B0 000110000? 001101102? 000??

Prophantis cf. androstigmata 01????11?0 00??01??11 10011????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????0 000110000? 0000?1105? 000?3

Table 2 continued.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

156

Character 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000000 0000000001 1111111111 11111

0000000001 1111111112 2222222223 3333333334 4444444445 5555555556 6666666667 7777777778 8888888889 9999999990 0000000001 11111

Taxon: 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 1234567890 12345

Prophantis xanthomeralis 01000?11?0 00000100?0 0001100000 0111010000 ?110011011 0000?00001 0001032101 0000100200 1000?101D? ?????????? ?????????? ?????

Pycnarmon pantherata 0100011001 0000000010 0001100000 0111010000 ?010210010 0000000100 ???1031100 0110000000 1001010122 000000000? 001101103? 000??

Rhectosemia multifarialis 0100010111 00?0000010 0001000000 0110010000 ?011011000 0000000010 ???0120100 0010000000 10010001B0 000110000? 0000?1103? 000??

Rhimphalea cf. astrigalis 0100011021 0000010010 1001100002 0111000001 0010010312 20000?1111 0101020100 0111000200 1001010100 000110000? 000100???? 000??

Salbia cf. haemorrhoidalis 01000?11?0 0000010??? 1001000000 0111010000 ?011211012 0000?31101 11000????0 10?1000200 0000?111D? 0001000010 001101100? 000?0

Samea cf. multiplicalis 0100111121 0000010010 00010????? ?????????? ?????????? ?????????? ?????????? ?????????? ?????????2 100110011? 1010?0???? 000??

Samea ecclesialis 0100111121 0000010010 0001100100 0011010001 0011111010 0000020001 0000120100 0011101201 1001010110 000100000? 000111101? 000??

Siga liris 0000000020 0000000010 0000?00000 000??00000 ?011211010 2100000010 ??00120100 0000000200 1000?10121 011110000? 00010110?? 000??

Spilomela perspicata 0100001021 0000011010 0001100000 010??00001 0011111321 0000000101 00010030?0 0001000100 1001010122 0000?0000? 0010?11100 000??

Spoladea recurvalis 0110001000 0000010010 0001100000 0111010001 0010011102 2100100001 1100120120 0000100210 1000?101D2 000010000? 0010?1A05? 000?0

Syllepis marialis 0100111111 0000010010 0000?00000 010??00100 ?01001101? 0000000000 ??000????0 00?0000000 10010001B2 000010000? 000101102? 0010?

Syngamia ﬂorella 0100001020 0000010010 1001100000 0111010001 00102110?? ??00001001 0000122100 0111000200 1000?1112? 0011100011 0000?1105? 000??

Terastia meticulosalis 0100111120 0100010010 1001101000 0111010001 001000??1? 4100100001 0001020101 0111100100 0000?10120 100110000? 000100???? 000?G

Trichaea pilicornis 0100101021 0000010010 1001100000 011001000? ?011011012 2100100001 0000122101 0010000200 0000?11120 000110000? 0010?1102? 000??

Udea ferrugalis 0100011100 0000010010 1??0?00000 010??00000 ?01101100? 0000100000 ??00120100 0010000000 10010001B2 0001000011 0000?1102? 000?0

Udea washingtonalis 0100011100 0000000001 1000?00000 000??10000 ?011011002 0000100010 ??00120100 0010000000 1000?01122 0001010011 100111102? 000??

Udeoides muscosalis 01000?11?0 00000100?0 1??1000000 000??10000 ?01101100? ??00110010 ??00120100 0020000000 1000?0011? ?????????? ?????????? ?????

Zebronia phenice 01000111?1 0100010010 0001000000 010??10000 ?011011020 0000?00001 0001020100 0111000201 1000?00112 0001000011 0000?0???? 000?0

Achyra cf. rantalis 1100011100 0000010111 01?1100000 0?10010000 ??10010002 3000200001 0000110100 0110000100 1000?011D0 0001000011 0000?11130 00100

Anania hortulata 0100001101 0010010110 11?1100000 0111010000 ?010010102 0010010000 ??00131110 0110000100 10011011D2 0011001010 ?00111103? 00100

Anania verbascalis 0100011120 0010010110 11?1100000 0111010000 ?010010102 0000010000 ??00110110 0110000100 10010011D2 000100100? 0000?1103? 00100

Cryptosara caritalis 0100111120 0000010110 11?1100000 2111010000 ?01100??02 3000200001 1001023100 0010000100 11011011D0 0001010011 100111103? 010??

Euclasta gigantalis 0100?11101 00?0000110 01?10????? ?????????? ??110?2?12 2100?00001 00000????0 00?0?00200 10011?01?0 0001000011 000111A06? 010??

Euclasta splendidalis 0100111121 0000000110 0??0?00000 0111010001 0011012012 ??00100001 00000????0 00?0000200 00010001D? 0101100011 1000?1106? 010??

Hyalobathra illectalis 0100111101 0000010111 11?1100000 0111010000 ??1110??0? 4100?00001 0000113100 000000??00 1101100120 0001?0000? 0000?1103? 010??

Hyalorista cf. taeniolalis 01001?11?0 00000101?1 01?1100000 0111010000 ?011011002 0000110001 0001010110 0000000100 1000?001D0 000110000? 000111103? 0010?

Loxostege aeruginalis 1100011121 0000010110 01?1000000 0111010000 ?010011012 1?00010010 ??00101100 0010000100 10010011D0 0011000010 000111103? 00100

Oenobotys sp. 01001?11?0 00100101?1 01?1100000 0?0??10000 ?010010102 0000210001 0000131110 0110000100 1000?001D? ?????????? ?????????? ?????

Ostrinia nubilalis 0100011100 0000010111 11?1100000 0011010000 ?010110002 3100000001 00001G0110 0111000100 00010001B2 0001?1001? 0000?1103? 0010D

Pagyda salvalis 0100001100 0000010111 11?1000000 0?11010000 ?01001100? 0010110001 0000120110 0020000000 10011001D0 100110000? 000111102? 0010?

Paracorsia repandalis 0100001100 00?0010011 01?1100000 0011010000 ?010010002 0000210011 0000120100 0011000100 10011001D? 000110000? 000111103? 00100

Portentomorpha xanthialis 0100111121 0000010011 11?1100000 0111010000 ?11100??20 3000100001 10001230?0 0000000000 1100?001D2 0001000011 001101102? 0011?

Psammotis pulveralis 0100111101 0000010111 01?1000000 0111010000 ?010010102 0000110000 ??00100110 0011000100 10011011D2 0011100011 0000?1103? 0010D

Pseudopyrausta cf. minima 0100001120 0000010111 01?1100000 0011010000 ?010010102 0000110000 ??00111110 0100000100 10010001D? ?????????? ?????????? ?????

Pyrausta purpuralis 0100011100 0000010111 01?1000000 0?0??10000 ??10011002 0000010000 ??01011100 0000000100 10010001D0 1000000011 0000?1103? 00100

Sitochroa verticalis 1100011120 0010010111 01?1000000 010??10000 ?010011002 1000100001 0000120100 0010000100 1000?011D2 001110011? 1000?1103? 00100

Tetridia vinacealis 0100011120 0000010010 01?1000010 0111010000 ?01001C012 ????001101 0010100100 0110010000 1001011102 0101000010 0001111161 ?10??

Uresiphita gilvata 0100111100 0010010111 11?0?00000 0111010000 ?010011012 ??00210001 0000101100 0111000000 1010?101B2 000110000? 000101102? 00100

Table 2 continued.

157

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46 Presence of chaetae on surface of uncus head(s): (0)

absent (Fig. 15C); (1) present (Fig. 3D).

47 Structure of uncus head chaetae: (0) simple, not api-

cally split (Figs. 7D, 14C); (1) bifurcate (Fig. 9I);

(2) multifurcate (popescu-gorJ & constantinescu

1977: g. 7d); (3) short, at, spatulate (Fig. 14C).

48 Location of setae on uncus: (0) dorsal (Fig. 13A);

(1) dorsal & lateral (Fig. 5E); (2) dorsal & ventral

(MallY & nuss 2010: g. 2B); (3) ventral (Fig. 9E);

(4) lateral (slaMka 2008: pl. 29 g. 182).

49 Attachment of uncus to tegumen: (0) broad, point of

attachment constricted (Fig. 4E); (1) broad, smooth

transition (Fig. 4F); (2) narrow, offset (Munroe

1976b: pl. u g. 6a; MallY & nuss 2010: g. 2C).

50 Region between subscaphium and dorsal tegumen:

(0) membranous (Fig. 9G); (1) sclerotized as gnathos

(sensu Maes 1998) (Fig. 3G); (2) sclerotized as pseu-

dognathos (sensu Maes 1998) (Figs. 4E, 5D).

51 Shape of transtillum arms: (0) triangular (tapering

towards apex) (Fig. 4E); (1) rounded (Fig. 3G); (2)

strap-like (apex blunt or pointed) (Figs. 8B, 11E);

(3) large rectangular, medioventrally with nger-like

process (“transtillum inferior” sensu Marion 1954)

(Fig. 15D; Marion 1954: g. 11); (4) rhomboidal (=

triangular with cut apex) (Fig. 15C).

52 Connection point of transtillum arms: (0) narrow

(Fig. 3G); (1) broad (Fig. 11E).

53 Presence of long dorsad chaetae on surface of tran-

stillum arms: (0) absent (Fig. 3G); (1) present (Fig.

15D).

54 Presence of lobar processes carrying hair-pencils

on the dorsolateral tegumen sides: (0) absent (Fig.

13B); (1) present (Fig. 3A).

55 Depth of gap/split of juxta: (0) < 10% of dorsoven-

tral length of juxta (Fig. 3G); (1) 10 – 60% of dorso-

ventral length of juxta (Fig. 5D); (2) > 60% of dorso -

ventral length of juxta to complete division into two

juxta arms (Figs. 3A, 6E).

56 Saccus shape: (0) U-shaped (Fig. 3A); (1) (sharply)

V-shaped (Fig. 4F); (2) stout, almost rectangular

(Fig. 5E); (3) narrow elongate (Fig. 9H).

57 Presence of constriction at basal saccus: (0) absent

(Fig. 3A); (1) present (Fig. 13A).

58 Ratio between saccus length and sacculus breadth:

(0) ≤ 1 (Fig. 3A); (1) > 1 (Fig. 13B).

59 Presence of protruding keel on ventral saccus tip: (0)

absent (Fig. 3A); (1) present (Fig. 4F).

60 Presence of partly sclerotized, chaetose hairpencil

articulating with the anterior edge of the vinculum-

tegumen connection: (0) absent (Fig. 7C); (1) pre-

sent (Figs. 13A, 10C, 14E).

61 Number of hairpencil sclerites on each side of the

genitalia: (0) one (Figs. 13A, 10C, 14E); (1) two or

more (articulated with each other via membranes)

(Fig. 9H).

62 Presence of more than one kind of hairpencil chae-

tae: (0) absent (Fig. 14E); (1) present (clarke 1986:

g. 34a; kiMura et al. 2002: gs. 1 – 4).

63 Presence of a pair of sclerotized, hair-studded hair-

pencils articulating with the anteromedian edge of

the saccus: (0) absent (Fig. 5E); (1) present (Fig.

3G).

64 Presence of bula emerging from central inner val-

va: (0) absent (Fig. 3D); (1) present (Fig. 4F).

65 Presence of bula emerging from dorsal valva base

near costa base: (0) absent (Fig. 3D); (1) present

(Figs. 4E, 10C).

66 General shape of bula: (0) broad triangular (Fig.

8B); (1) elongate triangular, at least twice as long

as broad (Fig. 4F); (2) elongate needle-like to claw-

shaped (Figs. 4E, 7D); (3) as long as broad, circular

to squarish (Fig. 12C).

67 General orientation of bula: (0) ventrally directed

towards sacculus or distal sacculus (Figs. 4E, 8B);

(1) directed towards ventral sacculus base (Fig. 7C);

(2) directed towards distal valva (Fig. 3A); (3) di-

rected dorsally, towards tegumen/uncus (Fig. 3G).

68 Presence of chaetae on bula surface: (0) absent

(Fig. 4E); (1) present (Figs. 7C, 15D).

69 Structure of apex of chaetae on bula surface: (0)

simple (Figs. 7C, 15D); (1) some simple, some mul-

tid (= editum of Pyraustinae) (Yang et al. 2012: up-

permost arrow in g. 7A – D); (2) spatulate (clarke

1986: g. 34a).

70 Presence of raised ridge running from basal to dorso-

distal sacculus: (0) absent (Fig. 5D); (1) present (Fig.

12D).

71 Presence of nger-like process studded with simple

chaetae emerging from central sacculus: (0) absent

(Fig. 5D); (1) present (Fig. 9H).

72 Presence of extension (process in some cases) at dor-

sodistal sacculus: (0) absent (Fig. 4E); (1) present

(Fig. 4F).

73 Spatial association of bula with dorsodistal saccu-

lus (or its extension): (0) distant (Fig. 4F); (1) closely

associated, overlapping (Fig. 4E); (2) bula and dor-

sodistal sacculus fused (Fig. 11E).

74 Presence of ination of basal costa: (0) absent (Fig.

4E); (1) present (Fig. 8B).

75 Joint of basal valva costa (with vinculum) extended

into an elongate, rod-shaped process: (0) absent (Fig.

4F); (1) present (Fig. 8B).

76 Presence of long, sometimes loosely arranged chae-

tae on surface of costal base: (0) absent (Fig. 4E); (1)

present (Figs. 3A, 5E).

77 Presence of a knee-like bend of 60 – 80° in the post-

basal costa: (0) absent (Fig. 4E); (1) present (Fig.

13B).

78 General shape of post-basal costa (not the entire dor-

sal valva edge): (0) concave (Fig. 3A); (1) straight

(Fig. 5E); (2) convex (Figs. 8B, 9F,G).

79 Presence of a setose dorsad process on the basal cos-

ta: (0) absent (Fig. 3A); (1) present (Fig. 11E).

80 General structure of distal costa: (0) tubular (Fig.

3A); (1) broadening (Fig. 12E).

81 Costa following the course of (= in alignment with)

the dorsal valva edge (all the way) into subapical

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

158

valva region: (0) absent (Fig. 9H); (1) present (Fig.

4E).

82 Presence of detached costa from valval area, the

costa protruding freely dorsad instead: (0) absent

(Fig. 7C); (1) present (Fig. 15C; Munroe 1976b:

pl. u g. 6a; sHaffer & Munroe 2007: gs. 130,

133).

83 Presence of a recess in the course of the ventral val-

va edge: (0) absent (Fig. 4F); (1) present (Fig. 7D).

84 Presence of a coecum on the phallus apodeme: (0)

absent (Figs. 8C, 12D); (1) present (Figs. 3B, 4G).

85 Length of phallus coecum relative to phallus apo-

deme length: (0) < 10% (Fig. 4G); (1) ≥ 10% (Fig.

3B).

86 Presence of reduction of phallus apodeme scleroti-

sation to a ventral, longitudinally sclerotized strip

(the rest of the apodeme being more or less membra-

nous): (0) absent (Figs. 4G, 15D); (1) present (Figs.

7D, 10C, 13F).

87 Presence of a distinct sclerite in the posterior phal-

lus apodeme: (0) absent (Fig. 13F); (1) present (Fig.

4G).

88 Presence of sclerotisation on surface of vesica: (0)

absent (Fig. 7F); (1) present (Fig. 3B).

89 Type of vesica sclerotisation: (0) single cornutus

(Fig. 3B); (1) multiple cornuti (Fig. 13F); (2) granu-

lated area (Fig. 3B).

90 Orientation of everted papillae anales: (0) postero-

ventrad (Fig. 3F); (1) ventrad (Fig. 4H); (2) poste-

riad (Fig. 5F).

91 Dorsal end of papillae anales larger than ventral

end: (0) absent (Fig. 3F); (1) present (Fig. 13E).

92 Ventral end of papillae anales larger than dorsal end:

(0) absent (Fig. 3F); (1) present (Fig. 14G).

93 Presence of a strongly sclerotized frame (= lamella

antevaginalis) around the ostium bursae: (0) absent

(Fig. 5F); (1) present (Fig. 3F).

94 Presence of strong sclerotisation in the antrum: (0)

absent, with antrum more or less membraneous

(Fig. 11H); (1) present (Fig. 4H).

95 Presence of a longitudinal membranous strip in the

antrum sclerotisation: (0) absent (Fig. 9J); (1) pre-

sent (Figs. 5F, 10D,E).

96 Presence of thickened mesocuticle in the antrum:

(0) absent (Fig. 11G); (1) present (Fig. 4H).

97 Presence of a cone-shaped central structure (Ana-

nia-type) in the antrum: (0) absent (Fig. 11G); (1)

present (Fig. 15F; tränkner et al. 2009: arrows in

gs. 18 – 21).

98 Presence of a lateral blind-end evagination (diver-

ticulum) in the colliculum: (0) absent (Fig. 13D); (1)

present (Fig. 13E).

99 Presence of a strongly sclerotised colliculum ante-

rior of antrum and posterior of attachment of ductus

seminalis: (0) absent (Fig. 11G); (1) present (Fig.

5F).

100 Presence of a longitudinal membranous strip in the

colliculum sclerotisation: (0) absent (Fig. 13D); (1)

present (Fig. 5F).

101 Presence of thickened mesocuticle in the collicu-

lum: (0) absent (Fig. 5F); (1) present (Fig. 14G).

102 Point of attachment of ductus seminalis to female

genital tract: (0) at posterior ductus bursae, at or

near colliculum (Fig. 5G); (1) at anterior ductus

bursae (Fig. 7E); (2) at corpus bursae (Fig. 12F).

103 Demarcation between corpus bursae and ductus

bursae: (0) distinct by narrow anterior ductus trans-

forming into wide corpus bursae (Fig. 3E); (1)

indistinct or absent by wide anterior ductus trans-

forming into equally wide corpus bursae, i.e. uent

transformation of d.b. to c.b.) (Figs. 3F, 10E).

104 Presence of sclerotisation in ductus bursae: (0) ab-

sent (Fig. 3E); (1) present (Fig. 5F).

105 Intensity of ductus bursae sclerotisation: (0) weak

(granulose texture) (Fig. 13E); (1) strong (Fig. 5F).

106 Presence of sclerotisation in corpus bursae: (0) ab-

sent (Fig. 9J); (1) present (Fig. 3F).

107 Structure of corpus bursae sclerotisation: (0) a gran-

ulose area (Figs. 11G – H); (1) one or more clearly

delimited sclerites (= signum, Pl. signa) (Fig. 3F).

108 Number of signa: (0) one (Fig. 3E); (1) two or more

(Fig. 3F).

109 Shape of anterior-most signum: (0) circular,

spinose, can be invaginated as a spine (Fig. 8K); (1)

longitudinal slim, strip-like (Fig. 13E); (2) elongate

rhombical to ovate (longitudinal axis longer than

transverse one) (Fig. 5F); (3) transverse rhombical

to cross-shaped (longitudinal axis shorter than or

equally long as transverse one) (Figs. 3E, 15F – G);

(4) patch of protruding teeth/spikes (Fig. 3F); (5)

transverse, smooth or dentate line or arch, with or

without central posteriad leg (if present, then sig-

num Y-shaped) (Figs. 10E, 11H); (6) broad, medi-

ally constricted, resembling puckered lips (Figs.

14G,H).

110 Shape of second signum (located posterior of rst

signum): (0) circular, spinose, can be invaginated

as a spine (Fig. 8K); (1) longitudinal slim, strip-like

(Fig. 5F); (2) elongate rhombical to ovate (sHaffer

& Munroe 2007: gs. 299, 300); (3) patch of pro-

truding teeth/spikes (Fig. 3F).

111 Presence of a third, slim, strip-like signum posterior

of the two anterior signa: (0) absent (Fig. 8J); (1)

present (Fig. 8K).

112 Presence of appendix bursae on anterior ductus bur-

sae: (0) absent (Fig. 8K); (1) present (Figs. 14G,

15G).

113 Presence of appendix bursae on corpus bursae: (0)

absent (Fig. 8K); (1) present (Figs. 3F, 15F).

114 Point of attachment of appendix bursae on corpus

bursae: (0) lateral (Figs. 3F, 15F); (1) posterior (Fig.

9J); (2) anterior (Fig. 5G).

115 Locality of larval feeding: (0) concealed in rolled/

spun leaves or in a web (leutHardt et al. 2010: g.

1; HaYden et al. 2017: g. 19); (1) on leaf/fruit sur-

face (upper/underside) (HaYden et al. 2013: Line-

odes fontella); (2) boring in stems and/or branches

(sourakov 2011: gs. 6A, 7A, 10B); (3) boring in

159

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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owers, pods and fruits (sourakov 2011: gs. 3,

4); (4) on/in roots (gentY & Mariau 1975: gs.

3 – 5, 7); (5) on dead and decaying plant matter

(MurpHY 1990: pl. 15 g. J).

3.3. Phylogenetic results

The investigation of the gene data with DAMBE5 showed

no signs for signicant substitution saturation in the

three different codon positions of COI, CAD, EF-1a and

GAPDH. Codon positions nt1 and nt2 of IDH and RpS5

showed no signicant saturation, whereas in nt3 of these

two genes, some substitution saturation was observed.

This low level of substitution saturation was accepted as

of minor inuence for the phylogenetic analysis of these

data, so that no codon position was removed from the

nal dataset. This decision was supported by trial analy-

ses of the dataset with nt3 removed from IDH and RpS5

which showed a very similar topology and comparable

node support.

In phylogenetic pre-analyses, several taxa had con-

spicuously unstable positions in the phylogeny. These

most problematic ‘rogue’ taxa were identied using

RogueNaRok and excluded from nal analyses. One ex-

ception was Niphopyralis chionesis, which we decided to

keep in the dataset despite its long terminal branch in the

phylogenetic results.

Differences in the coverage of morphological data

coding affect the performance of the phylogenetic analy-

ses: MrBayes analyses containing morphological data

for the outgroup taxa performed worse than analyses

that only comprised morphological data for Pyraustinae

and Spilomelinae and where outgroups were coded as

‘?’. When outgroup morphological data is included in

the analysis, the parallel MrBayes runs do not converge

properly and the effective sample size is low for a num-

ber of parameters. All phylogenetic results stated and

discussed below are therefore based on the datasets that

only comprise the morphological data for Spilomelinae,

Pyraustinae and Sufetula. The potential causes and impli-

cations of outgroup coding are elaborated in the Discus-

sion section.

The parallel runs of all MrBayes analyses converged

sufciently after 30 Mio. generations, and ESS were

(mostly well) above 100. The analyses of the different

datasets result in highly similar topologies, and branch

support from the analyses of the molecular data alone

and those of the combined molecular and morphological

data are almost identical (Fig. 1). The additional mor-

phological data in the analysis of the combined dataset

does not result in improved resolution or branch sup-

port as compared to the results of the molecular data-

set. Branching differences are found in the position of

the clade Spilomelini (see dotted arrow in Fig. 1), and

within the clade Margaroniini. Pyralidae, Pyralinae, and

Crambidae are each monophyletic with high branch sup-

ports. In the Crambidae outgroup, Sufetula (Lathroteli-

nae) is sister to the “CAMMSS Clade” sensu regier et

al. (2012), with a clade Crambus (Crambinae) + Eudonia

(Scopariinae) sister to the “Wet Habitat Clade” (sensu

regier et al. 2012) Clepsicosma + (Midila + Schoeno-

bius), the latter belonging to Acentropinae, Midilinae

and Schoenobiinae, respectively. Sister to the Crambidae

outgroup is the “PS Clade” (sensu regier et al. 2012)

of Pyraustinae and Spilomelinae. Both Pyraustinae and

Spilomelinae are highly supported (1 PP) monophyletic

and moderately-supported (0.93 – 0.95 PP) sister to each

other.

Within Pyraustinae, Tetridia is sister to all other taxa.

The two Euclasta species form a monophylum (Euclas-

tini) that is sister to the remainder of Pyraustinae. A clade

Uresiphita + (Portentomorpha + (Cryptosara + Hyalo-

bathra)) is sister to the remainder of Pyraustinae (Pyraus-

tini). Ostrinia is sister to a clade Pagyda + Paracorsia,

which is sister to the remainder of Pyraustinae. Achyra +

(Loxostege + Sitochroa) is sister to a clade Oenobotys +

(Hyalorista + Pyrausta) and its sister group of Psammo-

tis, Pseudopyrausta and Anania. Anania, with two sam-

pled species, is monophyletic except in the phylogram of

the GENES-partitioned genetic dataset, where A. hortu-

lata is sister to Psammotis, and A. verbascalis sister to

Pseudopyrausta, all with PP < 0.9.

The phylogenetic relationships within Spilomeli-

nae are as follows: Hydririni + ((Udeini + Lineodini) +

(Wurthiini + (Agroterini + (Margaroniini + (Spilomelini

+ (Herpetogrammatini + ((Hymeniini + Asciodini) +

(Trichaeini + (Steniini + Nomophilini))))))))), with the

exception of the GENES-partitioned analyses, where

Spilomelini is in an unsupported (0.67 – 0.72 PP) sister-

group relationship with Margaroniini (indicated by dotted

arrow in Fig. 1). Hydririni comprises Hydriris + (Lam-

prosema + (Gonocausta + Syllepis)). Udeini comprises

Conchylodes + (Udeoides + Udea). Lineodini comprises

Lineodes + (Rhectosemia + (Leucinodes + Neoleuci-

nodes)). Wurthiini comprises Apilocrocis + (Aristebu-

lea + Niphopyralis). Agroterini comprises Pycnarmon +

((Neoanalthes + (Aetholix + Agrotera)) + (Haritalodes

+ (Phostria + Patania))). Margaroniini forms a large

polytomy with several moderately to well-supported

monophyla, which are: Asturodes + Maruca; Omiodes;

Prenesta; Liopasia + (Agathodes + Terastia); Hodeber-

tia + (Antigastra + Zebronia); (Azochis + Conogethes) +

Ghesquierellana + Megastes); (Agrioglypta + Obtusipal-

pis) + (‘Dichocrocis’ cf. zebralis + Glyphodes). Addition-

al taxa in Margaroniini with unresolved or unsupported

(PP < 0.9) relationships are: Cydalima, Filodes, Rhim-

phalea, Diaphania, Palpita, Botyodes, Cadarena, Leu-

cochroma, ‘Nacoleia’ insolitalis. Spilomelini comprises

(Siga + Eporidia) + (Spilomela + (Salbia + (Marasmia +

Cnaphalocrocis))). Herpetogrammatini comprises Eur-

rhyparodes + (Herpetogramma + Hileithia). Hymeniini

comprises Hymenia + Spoladea. Asciodini comprises

Asciodes + Arthromastix. Trichaeini comprises Trichaea

+ Prophantis. Steniini comprises (Dolicharthria + Meta-

sia) + (Duponchelia + Anageshna). Nomophilini com-

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

160

prises (Syngamia + (Ategumia + (Bocchoris + (Diasemia

+ Diasemiopsis)))) + (Desmia + (Mecyna + (Samea +

Nomophila))).

Parsimony analysis resulted in three cladograms of

35,100 steps. The strict consensus (Fig. 2) has 35,153

steps. Condensing these cladograms with “collapse[” did

not lengthen them, so ltering with the command “best”

was not necessary. Niphopyralis groups with Sufetula.

The topology of the outgroup CAMMSS clade is differ-

ent than that in regier et al. (2012). Euclastini diverges

rst in Pyraustinae. The second-diverging clade is Por-

tentomorphini including Portentomorpha, Hyalobathra,

and Cry pto sara. The topology within Pyraustini is sub-

stantially different. Tetridia and Uresiphita are subordi-

nate in Pyraustini, sister to Pseudopyrausta and Ostrinia,

respectively.

Fig. 1.

161

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

The three unweighted cladograms differ in only two

clades: 1) whether Lineodes or Rhectosemia is the rst-

diverging genus of Lineodini, and 2) the topology (Ste ni-

ini + (Nomophilini + Trichaeini)) versus (Nomophilini +

(Steniini + Trichaeini), with Nomophilini in a reduced

sense including Syngamia but not the Ategumia clade.

Other differences are 1) the grouping of Desmia with

Trichaeini and 2) the Ategumia clade (with Bocchoris,

Diasemia and Diasemiopsis) consistently being sister to

the clade of the other three tribes (Steniini, Trichaeini,

and Nomophilini s.str.).

IterPCR (pol & escapa 2009) did not suggest any

taxa or characters to recode. Implied weighting with k-

parameter values of 9 through 13 found cladograms (not

shown) with Niphopyralis sister to Aristebulea principis

Munroe & Mutuura, 1968, but the topologies are oth-

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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

162

erwise similar to the equal-weights results. The lengths

range from 35,159 to 35,192 steps.

Among the genera with more than one sampled spe-

cies, Euclasta, Udea, Patania and Omiodes are mono-

phyletic. Marasmia, Prenesta and Samea are paraphyl-

etic, and Glyphodes and Dichocrocis are polyphyletic.

The morphological data were mapped with WinClada

on the Bayesian consensus (Fig. 1, synapomorphies not

shown) and the parsimony consensus (Fig. 2) using slow

optimization (= delayed transformation, DELTRAN).

The results are stated in the diagnoses of the clades in

the taxonomy section. Although consensus trees are typi-

cally longer than the shortest actual cladograms, the extra

steps did not occur along the particular clades that we are

interested in diagnosing.

4. Phylogenetic classification

In this section we focus on the taxonomic circumscrip-

tion of Spilomelinae and Pyraustinae and the clades

found therein. We state synapomorphies and / or charac-

ters derived from slow optimization in the “Synapomor-

Fig. 2. Strict consensus of three parsimony cladograms of 35,100 steps, with morphological characters mapped with slow optimization. —

Symbols: ● unique apomorphies; ○ homoplastic apomorphies.

163

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

phies” paragraphs. Representatives of all proposed tribes

are illustrated in Figs. 3 – 15. The morphological charac-

ters indicated in the gures do not necessarily represent

apomorphies for the respective tribe. A checklist of all

Spilomelinae and Pyraustinae genera that are placed in

tribes is given in the Appendix.

Fig. 2 – Continuation.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

164

4.1. Spilomelinae + Pyraustinae (PS) Clade

Synapomorphies. No unambiguous synapomorphies

were found, as most of the Crambidae outgroup taxa

were left uncoded. Under slow optimization, the fol-

lowing three characters are found: 36:1, U-shaped scle-

rotisation of lateral and anterior edge of male sternite 8

present (also present in Schoenobiinae: Schoenobius);

65:1, presence of bula emerging from dorsal valva base

near costa base; 106:1, sclerotisation in corpus bursae

present.

Description. The uncus has bid chaetae. These dis-

tinctive chaetae are present in many Pyraustinae as well

as in most Spilomelinae. Bid chaetae are lost in some

spilomeline groups which have normal, hair-like mono-

lament chaetae on the uncus instead (e.g. Conchylodes

genus group in Udeini, Agroterini, several Margaro-

niini).

The costa of the valva is straight to concave. All

investigated Pyraustinae as well as the non-euspilome -

line clades (Hydririni, Udeini, Lineodini, Wurthiini) ex-

hibit male genitalia with a straight or concave costa (we

only refer to the costa here, and not to the whole dor-

sal valva edge). Most other Spilomelinae have a convex

costa.

The gnathos is reduced to a transverse strap, laterally

fused to the tegumen, and usually without a central pro-

cess. This is the “pseudognathos” of Maes (1998a), which

solis & Metz (2011) homologized with the gnathos: the

structure is simply reduced and fused. A few Pyrausti-

nae and Spilomelinae do have a central process like that

in most other Crambidae, such as Munroeodes Amsel,

1957, Sarabotys Munroe, 1964, Phaedropsis Warren,

1890, Patania Moore, 1888, Syllepte amando (Cramer,

1779), Deuterophysa Warren, 1889 and Mimudea War -

ren, 1892. Munroe (1964) considered this process to be

primitive and indicative of relationship with Evergestini,

but our results indicate that such processes are secondar-

ily derived and homoplastic.

The phallus apodeme is evenly sclerotized. All inves-

tigated Pyraustinae (except Uresiphita) and the non-eu-

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-

ing the length of the phallus; this character is reversed in

several Spilomelinae in the euspilomelinae clades.

The signum is rhombiform. This distinctive signum

is a traditional character of Pyraustinae s.str. (Munroe

1976a). It is a single sclerite with two axes, a major and

minor one, and has short spines or granules. Apart from

Pyraustinae, this signum type is found in modied forms

in the non-euspilomelinae clades (see below). Lamprose-

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

transverse axis is nearly absent, and the whole is elongate

and zipper-shaped (Syllepis, Udea, Conchylodes, Rhec-

tosemia) to nearly circular (Choristostigma); we refer to

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

body shapes that likewise vary from nearly circular to

elongate.

The corpus bursae has an appendix bursae. An ap-

pendix is present in most investigated Pyraustinae and

in most Hydririni as well as in Conchylodes and Sisyrac-

era (Udeini). This character is absent in all other investi-

gated Spilomelinae except for several Margaroniini and

Eporidia (Spilomelini), where it might be a secondary

development.

Remarks. Immature stages of Pyraustinae and Spilo-

melinae have not been studied in a phylogenetic context,

and characters consistently separating the two groups are

not known (allYson 1981, 1984).

4.2. Spilomelinae Guenée, 1854

Type genus: Spilomela Guenée, 1854

= Sylleptinae Swinhoe, 1900

Synapomorphies. 10:1, maxillary palpi minute to ob-

solete, cannot hypothetically come in contact with each

other; 23:0, fornix tympani projecting in ventral direc-

tion (unique); 105:0, ductus bursae sclerotisation weak

or with granulose texture.

Description. The fornix tympani projects ventrad from

the tympanic frame. The retinacular hook (frenulum hook

sensu forbes 1926) is lost. Females have two frenular

bristles, while the number of female frenular bristles var-

ies in Pyraustinae.

Systematics. Spilomelinae includes a monophylum that

we refer to as “euspilomeline clade” (Greek eu- good,

true), characterised by two morphological synapo-

morphies (see below). In contrast, the tribes Hydririni,

Udeini, Lineodini and Wurthiini represent a paraphylum,

of which Wurthiini is sister to the euspilomeline clade.

Because of this paraphyly, we refrain from proposing a

name for the group, and refer to them as the non-euspi-

lomeline clades.

4.2.1. Non-euspilomeline clades

The non-euspilomeline clades are characterised by ple-

siomorphies shared with Pyraustinae: the pleural mem-

branes of the male abdominal segment 8 lack a longitu-

dinal sclerotized strip; the valva costa is straight or con-

cave; the phallus apodeme is evenly sclerotized; and the

signum is “ediacaroid”. Several taxa exhibit an appendix

bursae.

4.2.2. Hydririni + Lineodini

Synapomorphies. 9:1, intersexual size difference of 3rd

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.

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166

4.2.3. Hydririni Minet, 1982 stat.rev.

Type genus: Hydriris Meyrick, 1885

Synapomorphies.113:1, appendix bursae present on cor-

pus bursae (see remarks below). Most Hydririni exhibit

the unique apomorphy 38:1, central hair scale patch(es)

on anterior edge of male sternite 8 present. In the par-

simony trees, only 113:1 is an apomorphy of Hydririni.

Description. The genitalia morphology is heterogene-

ous: the valvas are slender to broad (Syllepis), and the

uncus and gnathos are reduced (Choristostigma War-

ren, 1892, Hydriris Meyrick, 1885, Hyperectis Meyrick,

1904) to well-developed. Choristostigma, Hydriris, Hy-

perectis, Nehydriris Munroe, 1974a and Rhectothyris

Warren, 1890 have a dorsolateral tegumen exhibiting

lobar processes with a eld of long, thin hair-pencils,

with a long phallus caecum, and with a single straight or

hooked cornutus. The hairs on the anterior edge of male

sternite 8 are absent in Choristostigma, Ommatospila

and Hydriris ornatalis but present in H. aonisalis. Some

taxa have an appendix bursae emerging laterally from the

corpus bursae as in Pyraustinae: Pyraustini (see below).

The signum is ediacaroid, circular to elongate with the

minor transverse axis varying from broad (Lamprosema

Hübner, 1823, Gonocausta Lederer, 1863) to short, or

forming circle(s) of radiating spines (Choristostigma,

Hydriris, Nehydriris). Ommatospila has a circular edi-

acaroid signum and an opposing signum consisting of a

eld of spines, like in Choristostigma and Hydriris.

Systematics. Minet (1982) established Hydririni in Gla-

phyriinae and included only Hydriris in this tribe. Mun-

roe (1995) returned Hydriris to Spilomelinae. Beside

Hydriris, Munroe (1995) placed in his Hydriris genus

group also Choristostigma, Geshna Dyar, 1906 and Ne-

hydriris.

According to our phylogenetic analysis, Hydriris (7

spp.), Gonocausta Lederer, 1863 (4 spp.), Lamprosema

(72 spp.) and Syllepis Poey, 1832 (7 spp.) belong to Hy-

dririni. Furthermore, based on morphological characters

we place Choristostigma (10 spp.), Nehydriris (1 sp.),

Ommatospila Lederer, 1963 (3 spp.) and Rhectothyris (1

sp.) here. According to morphological characters, Gesh-

na does not belong to Hydririni, but to Spilomelini (see

below).

Hyperectis dioctias Meyrick, 1904, the type species

of Hyperectis, is depicted in ziMMerMan (1958). From

there it is evident that this genus is misplaced in Pyraus-

tinae and that the genitalia are close to those of Hydriris

ornatalis (Duponchel, 1832) and H. aonisalis (Walker,

1859), and that the genus is not distinguishable from

Hydriris. We therefore synonymize Hyperectis Meyrick,

1904 syn.n. with Hydriris, and transfer the two species

Hydriris dioctias Meyrick, 1904 comb.n., and Hydriris

apicalis (Hampson, 1912) comb.n.

Food plants. The known larval food plants are Sapin-

daceae (Gonocausta, Lamprosema, Syllepis) and single

cases of Fabaceae (Lamprosema), Anacardiaceae and

Lamiaceae (both Syllepis) (Janzen & HallwacHs 2009).

Hydriris ornatalis larvae feed on the leaf undersides of

Ipomoea batatas (Convolvulaceae) and related plants,

later instars skeletonize the leaves (HinckleY 1964).

Remarks. The genitalia of Syllepis and Gonocausta are

highly similar, and a future revision might evaluate these

two genera as congeneric. Lamprosema contains numer-

ous misplaced Old-World species and needs revision. We

verify the congenerity of the taxon used in our analyses

with Lamprosema lunulalis Hübner, 1823 from Suri-

name, the type species of the genus.

An appendix bursae is also observed in Conchylodes

and Sisyracera (Udeini) as well as in Pyraustinae. Un-

der a slightly different basal branching sequence, the ap-

pendix bursae could be recovered as a synapomorphy of

Pyraustinae and Spilomelinae but lost in most Spilomeli-

nae. However, the best topology in this study indicates

separate origins.

Some of the characters by which Minet (1982) placed

Hydririni in Glaphyriinae are homoplastic. Spatulate

hind wing scales are paralleled with Glaphyriinae, and

solis & adaMski (1998) found that such scales are vari-

able even within Glaphyriinae. The spinose signa of H.

ornatalis resemble the spinose sclerotizations of many

Neotropical glaphyriines, but H. aonisalis has a lenticu-

lar ediacaroid signum.

4.2.4. Lineodini Amsel, 1956 stat.rev.

Type genus: Lineodes Guenée, 1854

Synapomorphies. 104:0, sclerotisation in ductus bursae

absent. Slow optimization only: 8:1, direction of 3rd labi-

al palpomere porrect (paralleled in other early-diverging

clades; not found in the parsimony trees); 95:1, longi-

tudinal membranous strip in the antrum sclerotisation

present. Fast optimization only: 19:0, female with only

one frenular bristle (HaYden et al. 2013); 106:0, signum

absent (not with slow optimization due to position of

Rhectosemia).

Description. The wings are moderately broad (Leuci-

nodes) to narrow and almost pterophorid-like in Lineodes

Guenée, 1854 and Atomopteryx Walsingham, 1891. The

sacci tympani are ventrally open (HaYden et al. 2013).

The valvae are very slender to relatively broad, trian-

gular or paddle-shaped, and the valva apex is rounded

to somewhat acute; the costa is straight to concave; the

bula is either slender and emerging from the costa base,

shorter and emerging more from the centre of the valva,

or entirely absent in Euleucinodes Capps, 1948 and Pro-

leucinodes Capps, 1948 (see capps 1948). The sacculus

is simple or (in Leucinodes Guenée, 1854) with a dis-

tal sacculus process in close association with the bula.

The posterior phallus is unmodied or with sclerotized

appendages (in Leucinodes, see MallY et al. 2015). The

posterior ductus bursae, colliculum and antrum in Leu-

cinodes and Neoleucinodes Capps, 1948 often have a

thickened mesocuticle and partial sclerotisation (HaYden

et al. 2013; MallY et al. 2015).

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Systematics. Lineodini was originally proposed for Li-

neodes (38 spp.) and Atomopteryx (10 spp.) (= Steno-

ptycha Zeller, 1863) (aMsel 1956); it is expanded here

to contain Leucinodes (20 spp.), Neoleucinodes (9 spp.)

and Rhectosemia Lederer, 1863 (12 spp.) according to

our phylogenetic analysis as well as to contain Euleuci-

nodes (1 sp.) and Proleucinodes (4 spp.), and to conrm

Atomopteryx based on morphological characters.

With the exception of Leucinodes, all these genera

were included by Munroe (1995) in his Udea genus

Fig. 4. Lineodini. A: ventral view of wings on frenulum bristle of female Leucinodes orbonalis. B: head of Le. orbonalis, male (left) and

female (right) (modied from Figs. 11 – 12 of MallY et al. 2015). C: adult male of  Lineodes vulnica. D: adult female of Le. Africensis.

E: male genitalia of Li. vulnica. 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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

168

group, plus Lamprosema and Udea, which however be-

long to Hydririni (see above) and Udeini, respectively

(see below).

The position of Rhectosemia in the phylogram (Fig. 1)

diverging after Lineodes disagrees with morphology. Fe-

males of some examined species of Rhectosemia have

two frenular bristles, and they have a signum that is usu-

ally narrow and elongate. These are plesiomorphies in

contrast to the synapomorphies of the other genera: one

bristle in both sexes and the loss of the signum.

Food plants. Almost all known larval food plants are

Solanaceae, the larvae either boring into the fruits (Leu-

cinodes, Neoleucinodes) or feeding on leaves and fruit

surfaces (Atomopteryx, Lineodes) (HaYden et al. 2013).

Several species of this group are pests on solanaceous

crops, e.g. Neoleucinodes elegantalis on tomato (Sola-

num lycopersicum), and Leucinodes spp. on eggplant

(Solanum melongena) (HaYden et al. 2013; MallY et al.

2015). Janzen & HallwacHs (2009) report two Neoleuci-

nodes species from Heliconia spp. (Heliconiaceae).

Remarks. Character 95:1, the presence of a longitu-

dinal membranous strip in the antrum sclerotisation,

is also present in Hydririni: Gonocausta, Syllepis. The

reduction to one frenular bristle in females is also pre-

sent in members of the Udea itysalis and U. alpinalis

species groups (sensu MallY & nuss 2011) (Udeini), in

Metasia suppandalis (Steniini), Diasemiopsis ramburi-

alis (Duponchel, 1833) (Diasemiini) and Niphopyralis

(Wurthiini). We nd Lineodini and Udeini (see below) to

be sister groups in the Bayesian analyses. However, they

do not share any synapomorphies with each other, also

not under slow optimization. Lineodini and Hydririni are

sister-groups in parsimony analysis, sharing 9:1, 68:0,

99:0, and 109:3.

4.2.5. Udeini Mally, Hayden, Neinhuis, Jordal &

Nuss trib.n.

Type genus: Udea Guenée, 1845 (in Duponchel)

Synapomorphies. 55:1, depth of gap/split of juxta being

10 – 60% of dorsoventral length of juxta; 99:1, strongly

sclerotised colliculum anterior of the antrum and poste-

rior of the attachment of the ductus seminalis present (not

found with parsimony); 109:2, signum elongate rhombi-

cal to ovate, longer than wide (found with parsimony).

Description. The uncus varies from unicapitate in the

Udea group (Deana Butler, 1879, Mnesictena Meyrick,

1884, Tanaophysa Warren, 1892, Udea Guenée in Du-

ponchel, 1845, Udeoides Maes, 2006) to conical (Con-

chylodes Guenée, 1854), reduced to triangle in Sisyrac-

era Möschler, 1890 and Ercta Walker, 1859, and reduced

to a transverse arching band in Cheverella Landry, 2011.

The uncus dorsally has bifurcate chaetae in the Udea

group, but the chaetae are simple and located dorsally

and ventrally in Conchylodes, Sisyracera and Cheverel-

la, and lost in Ercta. The costa of the valva is slightly

concave; the ventral sacculus edge is parallel to the costa

(inated in Cheverella), the valva apical of the sacculus

tapers towards a rounded apex. The female genitalia have

an elongate signum that is rhombical, lanceolate or edi-

acaroid in shape. In all Udea species groups sensu MallY

& nuss (2011) except the U. ferrugalis species group, an

accessory signum in the conjunction of ductus- and cor-

pus bursae is present. Conchylodes, Ercta and Sisyracera

have a membranous appendix bursae, attached anteriorly

in Conchylodes and Ercta and posteriorly in Sisyracera.

The antrum is strongly sclerotized, weakly in Cheverella.

Systematics. Minet (1982) associated Udea (214 spp.)

with Pyraustinae, a decision followed by leraut (1997,

2012). In contrast, our phylogenetic analysis supports

Udea as belonging to Spilomelinae, forming a monophy-

lum together with Udeoides (5 spp.) and Conchylodes

(21 spp.); furthermore, based on morphological charac-

ters, we place Cheverella (1 sp.), Deana (1 sp.), Ercta (7

spp.), Mnesictena Meyrick, 1884 (7 spp.), Sisyracera (3

spp.) and Tanaophysa (2 spp.) in this monophylum.

Udeini was proposed by leraut (1997) in Pyrausti-

nae, but without a description to differentiate the taxon,

a requirement by the International Code of Zoological

Nomenclature (ride et al. 1999, International Commis-

sion on Zoological Nomenclature: article 13.1) for names

published after 1930. Therefore, the family-group name

Udeini was not available prior to our proposal and formal

description.

The genitalia of Azochis graphialis Schaus, 1912,

type species of Nonazochis Amsel, 1956, resemble those

of C. diphteralis (Geyer, 1826), not justifying the sepa-

ration of the two genera. We therefore synonymize the

monotypic Nonazochis Amsel, 1956 syn.n. with Conchy-

lodes Guenée, 1854, and transfer Conchylodes graphialis

(Schaus, 1912) comb.n. Conchylodes octonalis (Zeller,

1873) comb.n. is transferred from Lygropia Lederer,

1863 based on characters in common with Conchylodes:

upward-curled transtilla arms, white wings with spots

(orange in C. octonalis, black in congeners), corpus bur-

sae with anterior appendix bursae, larvae feeding on Bor-

aginaceae (powell & opler 2009).

Food plants. The food plant spectrum is broad in Udeini,

and several Udea species such as U. ferrugalis, U. lutea-

lis, U. olivalis, U. prunalis and U. rubigalis are pronounc-

edly polyphagous (weigel et al. 1925; lHoMMe 1935).

Mnesictena  avidalis is recorded from Muehlenbeckia

(Polygonaceae), M. notata from Urtica and Australina

(Urticaceae) (robinson et al. 2010). The larvae of Con-

chylodes ovulalis (Guenée, 1854) are recorded to feed on

Platanus (Platanaceae) (solis 2008), other Conchylodes

species feed on Asteraceae, Cordiaceae, Malvaceae, Bor-

aginaceae and Annonaceae (Janzen & HallwacHs 2009).

Sisyracera and Cheverella are on Boraginaceae (dYar

1917; wolcott 1950; landrY et al. 2011).

Remarks. The genus Mnesictena was synonymised with

Udea by Munroe (1983), followed by sHaffer et al.

(1996). The type species of both genera were studied by

MallY & nuss (2011) and found to be not congeneric,

supporting dugdale’s (1988) view of keeping them as

separate groups, but the authors did not reinstate Mne-

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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sictena as bona genus. We leave this issue to a thorough

future study of the relationships within Udeini.

Sisyracera and Cheverella are problematic Neotropi-

cal genera. Munroe (1995) left Sisyracera unplaced, and

the relationship of Cheverella, a Galápagos endemic,

prompted lengthy discussion in landrY et al. (2011),

who decided that the Hydriris or Siga groups were the

most likely places. Their relationship with Conchylodes

Fig. 5. Udeini. A: adult female of Udea maderensis. B: adult male of Conchylodes ovulalis. 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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

170

seems more plausible to us. The larvae of all three genera

feed on Boraginaceae. The moths are white with black

spotted lines (dense reticulate pattern in Sisyracera) and

have ascending labial palpi. The valvae are attenuate, and

the uncus is reduced or lost, bearing only ne chaetae.

Sisyracera shares with Conchylodes an appendix bursae.

The homoplasies to be accounted for are the change of

the signum in Sisyracera and Cheverella (signum absent

in the former, a small thorn in the latter) and loss of the

valva bula in Sisyracera. The robust valvae and inated

sacculus of Cheverella can be explained as part of the

internally feeding larval syndrome, which is paralleled in

the Beebea group (Asciodini) and among the internally

feeding Margaroniini that had been classied in Mun-

roe’s (1995) polyphyletic Polygrammodes group.

Microphysetica Hampson, 1917 belongs to Hydririni

or Udeini. Females have a rhombiform signum and ap-

pendix bursae but not the tubular colliculum of Udeini.

Males have sternite 8 like Udea and Choristostigma:

centrally membranous but without a distinctive anterior

scale eld, and the juxta is mesally weak but not split into

distinct arms.

4.2.6. Wurthiini + euspilomeline clades

Synapomorphies. 109:0, shape of anterior-most signum:

circular, spinose, or invaginated as a spine, without obvi-

ous axes. 103:1, indistinct division of the ductus bursae

and corpus bursae (only found with parsimony).

4.2.7. Wurthiini Roepke, 1916 stat.rev.

Type genus: Wurthia Roepke, 1916 = Niphopyralis Hampson, 1893

Synapomorphies. 55:2, depth of gap/split of juxta rang-

ing from more than 60% of dorsoventral length of juxta

to complete division into two juxta arms; 66:1, general

shape of bula elongate, length at least twice the width,

apically rounded (may be curved). Slow optimization

only: 24:0, venulae secundae absent.

Description. The male genitalia have a broad triangu-

lar, ventrally directed bula; the mesal sides of the sac-

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 &

Mutuura, 1968); the mediodorsal sacculus has a medially

directed process (absent in Apilocrocis Amsel, 1956 and

Diaphantania Möschler, 1890). In the female genitalia,

the lamella antevaginalis forms a strongly sclerotised an-

trum frame; the signum is rounded, small (Aristebulea

Munroe & Mutuura, 1968, Pseudebulea Butler, 1881) to

relatively large (Apilocrocis, Diaphantania), and absent

in Mimetebulea and Niphopyralis. Under fast optimiza-

tion, the loss of venulae secundae (24:0) is shared with

various Hydririni, Lineodini and Udeini.

Systematics. Based on our phylogenetic results, we

place Apilocrocis (11 spp.), Aristebulea (2 spp.) and

Niphopyralis (= Wurthia) (8 spp.) in Wurthiini. Further-

more, based on morphological investigation, we place

Diaphantania (3 spp.), Mimetebulea (1 sp.) and Pseude-

bulea (4 spp.) in this tribe. This group can be considered

as an enlargement of Munroe’s (1995) Diaphantania ge-

nus group.

Food plants. Food plants are not known for most Wurthi-

ini. The larvae of Apilocrocis glaucosia (Hampson, 1912)

feed on Celtis iguanea (Ulmaceae) (Janzen & HallwacHs

2009). Niphopyralis larvae live as brood parasites in nests

of ants of the genera Oecophylla Smith, 1860 and Pol-

yrhachis Smith, 1857 and feed on eggs, larvae, and pupae

of their hosts (roepke 1916; keMner 1923).

Remarks. The placement of Niphopyralis in Spilomeli-

nae was a surprising discovery of regier et al. (2012),

but its particular association with Aristebulea and Api-

locrocis in our analysis allows a radical but satisfying

reinterpretation of the aberrant male genitalia (Fig. 6E).

The genitalia have been previously illustrated in Maes

(1998a), who interpreted the gnathos as consisting of two

separate, articulated arms. This condition occurs in other

lepidopteran superfamilies (e.g. Papilionoidea), but it is

not common in Pyraloidea (to our knowledge, occurring

elsewhere only in Heliothelinae). In our interpretation,

these two separate processes are the distal halves of the

true valvae. They are small and displaced dorsad, but they

have the same shape as the valvae in other wurthiines:

distally attenuate with a triangular swelling at the base of

the costa (like that in Diaphantania impulsalis (Herrich-

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

the outer margin in Apilocrocis and nearly so in the other

genera. We interpret the sclerotised structures anking

the juxta in Niphopyalis as the valva sacculi, each with a

median process similar to those in Aristebulea, Mimete-

bulea, Pseudebulea, Diaphantania and Apilocrocis. It is

not entirely clear whether the elongate processes in the

ventral region of the genitalic capsule arise from the sac-

culi or from the juxta (as coded in the character list 55:2).

We further interpret the pair of weakly setose structures

atop the tegumen to be the uncus in normal position,

even though the other members of the tribe have a single-

headed uncus with bid chaetae. The gnathos is absent.

4.2.8. Euspilomeline clades

Synapomorphies. 86:1, reduction of phallus apodeme to

a ventral, longitudinally sclerotized strip along the man-

ica (the rest of the apodeme being more or less membra-

nous). In addition, the parsimony trees add many more

synapomorphies: 32:1, male tergite 8 with anterior edge

emarginate; 60:1, partly sclerotized hair pencils present

on anterior edge of vinculum-tegumen connection; 78:2,

valva with convex costa; 84:0, phallus without coecum;

95:1, antrum with longitudinal membranous strip; and

99:0, a strongly sclerotised colliculum between antrum

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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

172

In this monophylum, the postmedial (PM) lines of

both wings are usually more jagged than in Pyraustinae

and among the early-diverging spilomeline clades. Es-

pecially the hindwing PM line is usually boldly marked

and projects distad on the M and CuA veins, whereas in

Pyraustinae, the hindwing PM line tends to be faint and

a smooth arc. In Pyraustinae, only very rarely does the

forewing PM line jut basad on the anal fold or is drawn

costad toward the discal spot (but see Pseudopyrausta).

In general, the hindwing PM line is similar to the fore-

wing PM line, so the combination of boldness and jag-

gedness distinguishes it. Although the jagged shape is

common in the euspilomeline clade, it is also present in

Aristebulea and Pseudebulea (Wurthiini).

4.2.9. Agroterini Acloque, 1897 stat.rev.

Type genus: Agrotera Schrank, 1802

Synapomorphies. 47:0, structure of uncus head chaetae

simple, not split; 58:1, ratio between saccus length and

sacculus breadth > 1 (elongate saccus, often with bul-

bous apex). The parsimony trees add 8:0, sclerotization

on vesica absent; and 44:0, conical (non-capitate) uncus.

Slow optimization only: 8:0, 3rd labial palpomere direct-

ed dorsally.

Description. The labial palps are upturned. The uncus

has a broad base, usually truncate to variously reduced,

at the extreme being a squat, transversely rectangular

square. The gnathos has a well-developed medial process

in some genera (see e.g. leraut 2005b: gs. 14 – 17).

The valvae are more or less rectangular, with costal and

ventral margins parallel, or slightly ovate; the saccus

is notably elongate, often distally bulbous. The female

genitalia have round and granular signa, single or double,

rarely extended as horns (Framinghamia Strand, 1920,

Phostria oajacalis (Walker, 1866)).

The upturned 3rd labial palpomere is a synapomor-

phy for this tribe in the DELTRAN analysis, but it is

shared with various other tribes, such as Asciodini, the

Siga group of Spilomelini, Spoladea, and some Nomo-

philini and Steniini. The presence of simple, unsplit

uncus chaetae is paralleled in some Pyraustinae, Marga-

roniini and Udeini, and in Nomophila Hübner, 1825. In

many genera, the tegumen mesally extends anteriad, like

an extended roof. This unique structure may characterize

a clade in Agroterini. In some genera, the papillae anales

face ventrad at a right angle to the axis of the ovipositor;

this state is paralleled in some Margaroniini.

Systematics. According to our phylogenetic results,

Agroterini comprises Aetholix Lederer, 1863 (4 spp.),

Agrotera (27 spp.), Haritalodes Warren, 1890 (11 spp.),

Neoanalthes Yamanaka & Kirpichnikova, 1993 (8 spp.),

Patania Moore, 1888 (= Pleuroptya Meyrick, 1890) (41

spp.), Phostria Hübner, 1819 (87 spp.) and ‘Pycnarmon’

pantherata Butler, 1878 which is not congeneric with

P. jaguaralis (Guenée, 1854), the type species of the

polyphyletic genus Pycnarmon Lederer, 1863 (59 spp.).

The placement of Pycnarmon among the euspilomeline

clades is still uncertain. Based on morphological charac-

teristics, we further place the following genera in Agro-

terini: Aiyura Munroe, 1974a (2 spp.), Bocchoropsis Am-

sel, 1956 (2 spp.), Chalcidoptera Butler, 1887 (15 spp.),

Chilochromopsis Munroe, 1964 (1 sp.), Coenostolopsis

Munroe, 1960 (3 spp.), Diastictis Hübner, 1818 (12 spp.),

Framinghamia (2 spp.), Glaucobotys Maes, 2008 (1 sp.),

Goliathodes Munroe, 1974a (1 sp.), Gypodes Munroe,

1976 (1 sp.), Lygropia Lederer, 1863 (68 spp.), Lypoti-

gris Hübner, 1825 (1 sp.), Micromartinia Amsel, 1957 (1

sp.), Microthyris Lederer, 1863 (7 spp.), Nagiella Mun-

roe, 1976 (4 spp.), Nosophora Lederer, 1863 (26 spp.),

Notarcha Meyrick, 1884 (18 spp.), Pantographa Leder-

er, 1863 (9 spp.), Phaedropsis Warren, 1890 (24 spp.),

Phryganodes Guenée, 1854 (26 spp.), Tetracona Mey-

rick, 1884 (2 spp.) and Ulopeza Zeller, 1852 (16 spp.).

Nagiella has been considered either a valid genus

(Munroe 1976b; kirti & sodHi 2001; rose 2001; ul-

laH et al. 2017) or a synonym of Pleuroptya (= Patania)

(inoue 1982; leraut 1997). We concur with Munroe’s

(1976b) separation of Nagiella from Patania. ullaH et

al. (2017) describe a fourth species in this genus. For the

generic diagnosis see Munroe (1976b).

This diverse, globally distributed tribe generally cor-

responds to Munroe’s (1995) Syllepte group. We conjec-

ture that he placed the Phaedropsis and Syllepte groups

rst in his checklist because some have a gnathos in the

traditional sense, i.e. with a well-developed medial pro-

cess, which would seem to be the primitive state. In our

analysis, this process is secondarily derived, since none

of the non-euspilomeline clades have it (the gnathos be-

ing a simple, transverse band). This process is also pre-

sent in two genera of uncertain placement: Mimudea

Warren, 1892, and Deuterophysa Warren, 1889.

Species of Phaedropsis are hardly separable from the

type species of Lygropia, Asopia unicoloralis Guenée,

1854. Lygropia and Phostria are major dustbin genera

of this tribe, holding many explictly misplaced species

(Munroe 1995).

In many genera, especially in the Old World, the fore-

wing costa bears a light-colored triangular spot. This is

the “Nosophora-Chalcidoptera” group referred to by

Munroe (1974a). In some taxa, the spot is so strongly

developed that it extends to the tornus and lls most of

the forewing (e.g. some misplaced ‘Leucinodes’ species,

‘Syllepte’ dottoalis Schaus, 1927).

Food plants. Larvae are generally leaf-tiers. Larvae of

Patania silicalis and P. sabinusalis have been reared

on Urticaceae (kiMball 1965; Miller et al. 2007; so-

lis 2008), P. silicalis furthermore on Polygonum (Po-

lygonaceae), Ipomoea and Merremia (Convolvulaceae),

Rivina (Petiveriaceae) and Bougainvillea (Nyctaginace-

ae) (Heppner & Habeck 1976; bendicHo-lopez 1998);

P. ruralis feeds on Urtica (Urticaceae), Humulus (Can-

nabaceae), Chenopodium, Atriplex (Amaranthaceae),

Filipendula (Rosaceae) and Ribes (Grossulariaceae)

(lHoMMe 1935); Central American Patania species (as

Pleuroptya) are recorded from Acanthaceae, Rubiaceae

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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and Urticaceae (Janzen & HallwacHs 2009). Known

food plants for Phostria larvae are mainly Convolvu-

laceae, Malvaceae and Rubiaceae (Janzen & HallwacHs

2009). Agrotera nemoralis feeds on Carpinus, Betula,

Corylus (Betulaceae), Castanea and Quercus (Fagaceae)

(Melzer & nuss 2009), while two Australasian Agrotera

species are reported to feed on Syzygium spp. (Myrta-

ceae) (Miller et al. 2007). Haritalodes is recorded from

Malvaceae, Amaranthaceae and Moraceae (gHesquiére

1942; Miller et al. 2007); Diastictis on Asteraceae (pow-

ell & opler 2009); Framinghamia on Salix (Salicaceae);

Phaedropsis on Polygonaceae and Malvaceae (Janzen &

HallwacHs 2009).

Signicant host associations are with Malvaceae

s.l. (Pantographa, Haritalodes, Phaedropsis), Convol-

vulaceae (Phostria tedea-group, Lygropia tripunctata-

group, Microthyris incl. Cyclocena; see HaYden & dickel

2014) and Rubiaceae (‘Pilocrocis’ xanthozonalis-group).

Remarks. The mimetic ‘Pilocrocis’ xanthozonalis-group

belongs here, and its species are misplaced in the Her pe-

togrammatini genus Pilocrocis.

The Australian species of Agrotera have recently been

revised by cHen et al. (2017), who removed Leucinodella

Strand, 1918, Nistra Walker, 1859, Sagariphora Mey-

rick, 1894 and Tetracona from synonymy with Agrotera;

the former three genera can currently not be placed in any

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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

174

of the proposed tribes, but Tetracona is placed in Agro-

terini, close to Aetholix (cHen et al. 2017).

4.2.10. Margaroniini Swinhoe & Cotes, 1889

stat.rev.

Type genus: Margaronia Hübner, 1825

= Dichocrociinae Swinhoe, 1900: 478

= Hapaliadae Swinhoe, 1890: 268

= Margarodidae Guenée, 1854: 286

Synapomorphies. 33:1, sclerotization of male tergite

8: heterogenous, i.e. with distinct sclerotization pattern;

72:1, extension (process in some cases) at dorsodistal

sacculus present (not found with parsimony); 75:1, joint

of basal valva costa (with vinculum) extended into an

elongate, rod-shaped process present; 103:0, demarca-

tion between corpus bursae and ductus bursae distinct,

with a narrow anterior ductus transforming into a wide

corpus bursae (only found with parsimony); 108:1, two

signa. Slow optimization only: 40:1, sclerite on each

pleural membrane of male segment 8 present (not found

with parsimony); 95:1, longitudinal membranous strip in

the antrum sclerotisation present (not found with parsi-

mony).

Description. The adult moths are mostly medium-sized

to large. Many genera have the scape and pedicel of the

male antenna modied. Males of many genera have a

large, black tuft of ne, hairlike sex-scales on A8. The

tegumen is often spacious in sagittal dimension (hamper-

ing the planar mounting of the genitalia on a glass slide).

The uncus is conical or unicapitate and long-necked (bi-

furcate in Cydalima perspectalis), with simple or bifur-

cate chaetae, often with both, or without chaetae. The

anterior tegumen-vinculum connection usually has an at-

tached pad of hairpencils, the pad structure ranging from

a simple sclerotized base with one kind of simple hairs to

several membrane-connected sclerites with several dif-

ferently structured hairs. The valva is commonly broad

and oval, with one bula about halfway to the apex. The

sacculus is comma-shaped, broadest at the anteroventral

valva base, arching and tapering distad (broadening in

Liopasia and Obtusipalpis), its distal apex usually ending

in a ridge or sclerotized process in close spatial associa-

tion with the distal bula. The vesica of the phallus has

a granulated area and/or one to several bodkin-shaped

cornuti. The corpus bursae often has a pair of circular

signa, which can be at or invaginated to form spikes,

or signum absent. The ductus bursae is granular in many

genera (e.g. Cydalima).

The heterogenous sclerotization of male tergite 8

(33:0) may be shared also with Asciodini, Spoladea,

Trichaeini, and some Steniini.

The base of the valva costa simple, not rod-shaped, in

several genera: the stout-bodied Liopasia Möschler, 1882,

Megastes Guenée, 1854 and Obtusipalpis Hampson,

1896, and in ‘Glyphodes’ rubrocinctalis (Guenée, 1854)

and Zebronia phenice (Stoll, 1782). The rod-shaped state

is paralleled in Hymeniini, Arthromastix lauralis (Walker,

1859), Samea ecclesialis Guenée, 1854 and Prophantis

xanthomeralis Hampson, 1918 comb.n.).

Systematics. Based on our phylogenetic ndings we as-

sociate these taxa with Margaroniini: Agathodes Guenée,

1854 (16 spp.), Agrioglypta Meyrick, 1932 (11 spp.),

Antigastra Lederer, 1863 (2 spp.), Asturodes Amsel,

1956 (1 sp.), Azochis Walker, 1859 (16 spp.), Botyodes

Guenée, 1854 (10 spp.), Cadarena Moore, 1886 (1 sp.),

Conogethes pandamalis (Walker, 1859) comb.n., Cyda-

lima Lederer, 1863 (9 spp.), Diaphania Hübner, 1818 (95

spp.), Dichocrocis cf. zebralis (Moore, 1867), Filodes

Guenée, 1854 (16 spp.), Ghesquierellana Berger, 1955

(5 spp.), Glyphodes Guenée, 1854 (156 spp.), Hodeber-

tia Leraut, 2003 (1 sp.), Liopasia (15 spp.), Leucochroma

Guenée, 1854 (6 spp.), Maruca Walker, 1859 (4 spp.),

Megastes (16 spp.), ‘Nacoleia’ insolitalis (Walker, 1862),

Obtusipalpis (6 spp.), Omiodes Guenée, 1854 (98 spp.),

Palpita Hübner, 1808 (162 spp.), Prenesta Snellen, 1875

(18 spp.), Pygospila Guenée, 1854 (10 spp.), Rhimphalea

Lederer, 1863 (12 spp.), Terastia Guenée, 1854 (7 spp.),

Zebronia Hübner, 1821 (6 spp.). ‘Nacoleia’ insolitalis is

misplaced in Nacoleia Walker, 1859, and its correct ge-

neric afliation remains uncertain. ‘Dichocrocis’ panda-

malis is misplaced in Dichocrocis; its correct placement

is in Conogethes Meyrick, 1884, where it is transferred

here (see above). Dichocrocis Lederer, 1863 (53 spp.) is

considered polyphyletic and needs revision; maculation

and male genitalia of the type species D. frenatalis Leder-

er, 1863 indicate a placement among the euspilomeline

groups, probably near or in Steniini, but this needs further

investigation.

‘Glyphodes’ rubrocinctalis is misplaced in Glyphodes;

in our phylogenetic analysis (Fig. 1) it is subordinate in

Prenesta. The male genitalia are smaller than those of

the type species of Prenesta, P. scyllalis, but they are

similar in structure, and the moths share distinctive red

and yellow maculation. We therefore transfer Prenesta

rubrocinctalis (Guenée, 1854) comb.n. from Glyphodes.

Furthermore, we assign the following taxa based

on morphological investigation: Alytana J.C. Shaffer

& Munroe, 2007 (2 sp.), Anarmodia Lederer, 1863 (24

spp.), Aphytoceros Meyrick, 1884 (3 spp.), Arthroschis-

ta Hampson, 1893 (2 spp.), Caprinia Walker, 1859 (11

spp.), Chabulina J.C. Shaffer, & Munroe, 2007 (2 spp.),

Charitoprepes Warren, 1896 (2 sp.), Chrysophyllis Mey-

rick, 1934 (1 sp.), Chrysothyridia Munroe, 1967 (2 spp.),

Cirrhochrista Lederer, 1863 (38 spp.), Colomychus

Munroe, 1956 (2 spp.), Compacta Amsel, 1956 (4 spp.),

Condylorrhiza Lederer, 1863 (4 spp.), Conogethes (16

spp.), Didymostoma Warren, 1892 (2 spp.), Dysallacta

Lederer, 1863 (3 spp.), Endocrossis Meyrick, 1889 (4

spp.), Eusabena Snellen, 1901 (4 spp.), Glyphodella J.C.

Shaffer, & Munroe, 2007 (3 spp.), Hedyleptopsis Mun-

roe, 1960 (1 sp.), Heterocnephes Lederer, 1863 (4 spp.),

Hoterodes Guenée, 1854 (5 spp.), Loxmaionia Schaus,

1913 (1 sp.), Marwitzia Gaede, 1917 (3 spp.), Mega-

physa Guenée, 1854 (1 sp.), Meroctena Lederer, 1863 (4

spp.), Nolckenia Snellen, 1875 (1 sp.), Omphisa Moore,

1886 (10 spp.), Pachynoa Lederer, 1863 (12 spp.), Paro-

175

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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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. rudiscalis. 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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

176

tis Hübner, 1831 (37 spp.), Poliobotys J.C. Shaffer &

Munroe, 2007 (1 sp.), Polygrammodes Guenée, 1854 (78

spp.), Polygrammopsis Munroe, 1960 (1 sp.), Radessa

Munroe, 1977 (2 spp.), Rhagoba Moore, 1888 (2 spp.),

Sinomphisa Munroe, 1958 (3 spp.), Sparagmia Guenée,

1854 (1 sp.), Stemorrhages Lederer, 1863 (8 spp.), Syn-

clera Lederer, 1863 (13 spp.), Syngamilyta Strand, 1920

(5 spp.), Talanga Moore, 1885 (9 spp.), Tessema J.F.G.

Clarke, 1986 (1 sp.), Tyspanodes Warren, 1891 (20 spp.),

Uncobotyodes Kirti & Rose, 1990 (1 sp.).

Tyspanodes is not a natural group, and at least T. ex-

althealis (Walker, 1859) is misplaced here; we did not in-

vestigate the type species, T. nigrolinealis (Moore, 1867),

but we can condently place T. hillalis Schaus, 1927, T.

hypsalis Warren, 1891 and T. celebensis Munroe, 1960 in

Margaroniini.

Heterocnephes apicipicta Inoue, 1963 is misplaced

in Heterocnephes and transferred to the monotypic

Charitoprepes as Charitoprepes apicipicta (Inoue,

1963) comb.n. The type species C. lubricosa Warren,

1896 shares with C. apicipicta the wing pattern and the

structure of the male genitalia (cf. inoue 1963; kiM et al.

2014). Alytana calligrammalis (Mabille, 1879) comb.n.

is transferred from Analyta, a transfer that had not been

formerly proposed by sHaffer & Munroe (2007) in their

description of Alytana.

Food plants. We arrange food plant records, where avail-

able, according to the clades found within Margaroniini

as shown in Fig. 1: Astrodes  mbriauralis is recorded

from Colubrina (Rhamnaceae); Maruca vitrata is a pest

species on various Fabaceae such as Lablab, Phaseolus,

Pisum, Psophocarpus, Sesbania and Vigna, but has also

been recorded from Rubiaceae, Solanaceae, Poaceae and

Euphorbiaceae (robinson et al. 2010). Cydalima mainly

feeds on Apocynaceae (C. laticostalis (Guenée, 1854)),

Buxaceae (C. perspectalis (Walker, 1859)) or Rhamnace-

ae (C. mysteris Meyrick, 1886) (robinson et al. 2010).

Filodes feeds on Thunbergia (Acanthaceae) (robinson

et al. 2010). Diaphania species mostly feed on Cucur-

bitaceae; Palpita is primarily on Oleaceae, but P. egia,

probably the rst-diverging member with a plesiomor-

phic male antennal scape, feeds on Apocynaceae (Moore

1884 – 1887; HinckleY 1964; kiMball 1965; claviJo al-

bertos 1990; solis 2006, 2008; robinson et al. 2010).

Omiodes species feed on a variety of host plants, and two

species (O. diemenalis (Guenée, 1854), O. indicata (Fab-

ricius, 1775)) are widespread pests on Fabaceae; the lar-

vae of the Hawaiian Omiodes clade feed on monocotyle-

donous plants, except O. monogona Moore, 1888, which

feeds on Fabaceae (robinson et al. 2010; Haines & ru-

binoff 2012). Omiodes stigmosalis Warren, 1892, a borer

in g fruits (Janzen & HallwacHs 2009), is misplaced in

Omiodes but has the characters of Margaroniini. Pren-

esta is recorded mainly from Apocynaceae and Moraceae

(Janzen & HallwacHs 2009). Larvae of Liopasia, Ag-

athodes and Terastia commonly feed on Erythrina (Fa-

baceae) (HinckleY 1864; kiMball 1965; sourakov 2012;

pereira et al. 2014). Antigastra catalaunalis (Duponche-

lia, 1833) and Zebronia phenice (Stoll in Cramer & Stoll,

1782) are leaf-tiers on Lamiales: The former, best known

as a pest of sesame (Pedaliaceae), also feeds on Bigno-

niaceae (Tecoma stans) and Plantaginaceae (powell &

opler 2009). The latter feeds on Bignoniaceae but was

also recorded on Gossypium (Malvaceae) and Ricinus

(Euphorbiaceae) (robinson et al. 2010). Hodebertia

testalis (Fabricius, 1794) larvae predominantly feed on

Asclepiadaceae (robinson et al. 2010). Botyodes feeds

on Flacourtiaceae, Moraceae, Salix (Salicaceae) and sev-

eral other hosts (nakaMura & oHgusHi 2004; robinson

et al. 2010). Cadarena pudoraria (Hübner, 1825) and

the closely related ‘Glyphodes’ (or ‘Pyrausta’) perel-

egans (Hampson, 1898) group are recorded from Passi-

oraceae, C. pudoraria also from Gossypium and Sida

(Malvaceae) (Janzen & HallwacHs 2009; robinson et al.

2010; de prins & Mazzei 2016). Ghesquierellana hirtu-

salis (Walker, 1859) larvae feed on Ficus (Moraceae) and

Gossypium (Malvaceae); Megastes on Ipomoea (Convol-

vulaceae); Azochis on Ficus (Moraceae); Conogethes lar-

vae are recorded from a wide range of plants, e.g. Pinace-

ae, Gnetaceae, Malvaceae, Sapindaceae, Euphorbiaceae

and Zingiberaceae (robinson et al. 2010; sHasHank et al.

2018). ‘Nacoleia’ insolitalis from Sandoricum (Meliace-

ae) (robinson et al. 2010). The known larval host plants

of the Glyphodes genus group sensu sutrisno (2002b)

(Glyphodes, Dysallacta, Talanga, Agrioglypta) are pri-

marily the latex-containing Moraceae and Apocynaceae

(kiMball 1965; coMMon 1990; robinson et al. 2010).

Obtusipalpis is recorded from Rubiaceae, Moraceae and

Rutaceae (robinson et al. 2010).

Host plants for other genera placed in Margaroniini

are: Arthroschista and Parotis on Rubiaceae, the latter

also on Apocynaceae, on which Pygospila and Stemor-

rhages mainly feed; Cirrhochrista on Moraceae; Con-

dylorrhiza on Salicaceae; Eusabena on Hoya (Asclepia-

daceae); Anarmodia and Sparagmia on Araliaceae; Syn-

clera on Gouania and Zizyphus (Rhamnaceae) (Mann &

brar 1980; Janzen & HallwacHs 2009; robinson et al.

2010; HaYden et al. 2017).

The most general trend in this group is feeding on la-

tex-bearing plants, especially Apocynaceae and Morace-

ae. The habit of boring in tubers of Ipomoea (Convol-

vulaceae) by Megastes, Polygrammodes eleuata, and

Omphisa anastomosalis (Guenée, 1854) is explained by

the presence of latex in these roots. Nevertheless, many

genera and genus groups diverge from the pattern and

radiate on non-latex-bearing plants, e.g. some Polygram-

modes in roots of Vernonia, Sinomphisa in Bignoniaceae,

and Omphisa fuscidentalis (Hampson, 1896) in bamboo.

Remarks. Margaroniini roughly reects a combination

of Munroe’s (1995) Diaphania and Polygrammodes

groups. The assumed close relationship for Agathodes,

Terastia and Liopasia (Munroe 1960; sourakov et al.

2015) has been conrmed by our results. Females of the

Agathodes genus group exhibit an appendix bursae (ab-

sent in some species); the larvae are feeding on Erythrina

(Fabaceae) (pereira et al. 2014).

The possible sister group relationship between Omi-

odes and Cnaphalocrocis Lederer, 1863, as suggested by

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

Haines & rubinoff (2012), could not be conrmed. We

treat Cnaphalocrocis as a member of Spilomelini (see

below), whereas Omiodes belongs to Margaroniini. The

sister group of Omiodes remains to be discovered.

claviJo albertos (1990) observed and described the

“anepisternal scale organ” in males of many genera of

Spilomelinae. Apparently, this is a tymbal organ (naka-

no et al. 2012b). Among Spilomelinae, ultrasound pro-

duction is reported from the Margaroniini Conogethes

punctiferalis (Guenée, 1854), Glyphodes pyloalis Walk-

er, 1859 and Palpita nigropunctalis (Bremer, 1864) as

well as in Spoladea recurvalis (Fabricius, 1775) (Hyme-

niini) (nakano et al. 2009, 2012a).

4.2.11. Spilomelini Guenée, 1854 stat.rev.

Type genus: Spilomela Guenée, 1854

= Siginae Hampson, 1918

Synapomorphies. 45:2, apical uncus bifurcate. The un-

cus is bicapitate with bulbous heads.

Description. Small (15 mm wingspan) to large (90 mm

wingspan) moths. Spilomelini shares the bicapitate uncus

with Asciodini and some Steniini such as Metasia and

Loxostegopsis. The costa of the valva is straight to slight-

ly concave or convex. This tribe consists of two distinct

clades: the Cnaphalocrocis group and the Siga group.

In the Cnaphalocrocis group, adults are small to

medium-sized. Most genera are brown and drab in col-

our, whereas Spilomela has contrasting maculation;

the wings’ transverse lines consist of straight segments

and angulate junctions. Males have a eld of enlarged,

raised scales on the centre of the forewing costa (absent

in several taxa). Sacci tympani are small and closely set.

In the male genitalia, the attened uncus is weakly bid,

the head consisting of two connected, at pads or elds

of chaetae rather than clearly separate parts; the uncus is

lost in Geshna. The sacculus ends distally with a small

bula-like process pointing inward toward the center of

the valva. Some taxa have a bula in the center of the

valva. In Spilomela perspicata, the details of the male

genitalia are distorted by elongation, but the uncus is

apically bid on close inspection. In females, the col-

liculum is cylindrical and open dorsally or entire. The

colliculum is often extended as extra sclerotization on

the adjacent ductus bursae. The ductus bursae is usually

very short and has ne spinules or striations next to the

colliculum. The signum typically is a granulose circle

or a small thorn, but Palpusia species have two long,

sickle-shaped signa, and Spilomela receptalis (Walker,

1859) has two shorter sickles; signa are absent in Rhec-

tocraspeda and Spilomela perspicata (Fabricius, 1787)

itself, which in addition has a very long, unsclerotized

ductus bursae.

The Siga group includes medium-sized to large and

thick-bodied moths. The proboscis is lost in Siga, other-

wise normally developed. The sacci tympani are exposed

as a shallow zona glabra, and the fornix tympani is circu-

larly rounded without an angle. The male genitalia have

the uncus entirely split into two separate unci (unsplit in

Zeuzerobotys), bearing bid chaetae; the costal margin of

the valva is approximately straight or only slightly con-

vex near the base, never strongly convex, distally straight

or slightly concave; the apical half of the valva is bluntly

attenuate, slightly to markedly narrower than basal half

of valva with its inated sacculus; there is one ventrally

directed, hook-shaped to spatulate bula emerging from

centre of valva. The combination of the bid uncus and

the shape of the valva distinguish members of the Siga

group from robust-bodied Margaroniini. The ductus bur-

sae is as long as or shorter than corpus bursae, and the

corpus bursae is spherical, rarely ovate, without signa.

Systematics. Based on our phylogenetic analyses, we

place the following taxa in Spilomelini: Cnaphalocrocis

group with Cnaphalocrocis (27 spp.), Marasmia Lederer,

1863 (9 spp.), Salbia Guenée, 1854 (35 spp.) and Spilo-

mela Guenée, 1854 (8 spp.); Siga group with Eporidia

Walker, 1859 (1 sp.) and Siga Hübner, 1820 (2 spp.), as

presumed by Munroe (1958).

Based on morphological investigation, we further as-

sign Aethaloessa Lederer, 1863 (3 spp.), Geshna Dyar,

1906 (1 sp.), Marasmianympha Munroe, 1991 (1 sp.),

Orphanostigma Warren, 1890 (6 spp.), Palpusia Am-

sel, 1956 (10 spp.) and Rhectocraspeda Warren, 1892 (2

spp.) to the Cnaphalocrocis group, and Cirrhocephalina

Munroe, 1995 (5 spp.), Scaptesylodes Munroe, 1976 (2

spp.) and Zeuzerobotys Munroe, 1963 (1 sp.) to the Siga

group, following Munroe (1963; 1976b; 1995) and lan-

drY et al. (2011) (but see Remarks).

Food plants. The hosts of the Cnaphalocrocis group are

heterogeneous, but the group includes a major radiation

on monocots, especially on Gramineae. The larvae of

Aethaloessa, Cnaphalocrocis, Marasmia and Salbia are

mainly leaf-rollers on Poaceae, Salbia larvae are also re-

corded from Verbenaceae and to a lesser amount from

Gesneriaceae and Fabaceae, and Aethaloessa  oridalis

(Zeller, 1852) from Urticaceae (Janzen & HallwacHs

2009; robinson et al. 2010). Rhectocraspeda is found

on Piper (Piperaceae), Columnea (Gesneriaceae) and

Solanaceae, Geshna on Canna (Cannaceae), Lilium (Lil-

iaceae), Thalia geniculata (Marantaceae) and Zantede-

schia (Araceae), Palpusia on Convolvulaceae and Ru-

biaceae, and Orphanostigma on Lamiaceae, Asteraceae

and Malvaceae (kiMball 1965; Heppner 2003; Janzen &

HallwacHs 2009; robinson et al. 2010). Spilomela larvae

are reported from Dilleniaceae, Ulmaceae and Rubiaceae

(Janzen & HallwacHs 2009).

In the Siga group, the hosts are unknown for Siga and

Eporidia, the two genera included in our phylogenetic

analysis.

Remarks. The genus Spilomela is polyphyletic and

needs revision. We base our conclusions on the type spe-

cies, S. perspicata.

The Siga group shares with Asciodini a similar mor-

phology of the male genitalia, especially the overall ro-

bust form and bid uncus. We transfer several genera to

Asciodini (below). Together with Siga and Eporidia, the

genera that we retain in the Siga group on morphologi-

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

178

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.

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cal grounds (Cirrhocephalina, Scaptesylodes, and Zeu-

zerobotys) share the same shape of bula and valva, and

non-inated transtilla. The colliculum of Siga, Eporidia,

and Scaptesylodes is entire and bulges ventrad rather like

a pot-belly or a pitcher plant (Nepenthes) (female genita-

lia not observed for Cirrhocephalina and Zeuzerobotys).

The loss of the proboscis in Siga is paralleled in Wurthi-

ini: Niphopyralis.

The monotypic Gesha is included because the larvae

feed on Canna, the wing pattern is typical of the Cnaph-

alocrocis group, and the genitalia share characters with

some Salbia species, namely swellings on the base of the

valva costa (also in S. mizaralis (Druce, 1899)) and the

broad, triangular uncus. The loss of bid uncus chaetae

and the movement of the bula to a central position on

the valva are homoplasies.

4.2.12. Herpetogrammatini Mally, Hayden,

Neinhuis, Jordal & Nuss trib.n.

Type genus: Herpetogramma Lederer, 1863

Synapomorphies. 5:1, length of sensillar setae at basal

antennomeres relative to diameter of basal antennomeres

in male > 50%; 107:0, corpus bursae sclerotisation con-

sisting of a granulose area. Slow optimization only: 8:1,

3rd labial palpomere porrect. The parsimony trees im-

ply one apomorphy: 44:0, a conical, non-captitate uncus;

characters 5:1 and 107:0 are synapomorphies with As-

ciodini.

Description. The uncus is conical, non-capitate, slen-

der to broad, with dorsally attached chaetae that are

bid in Eurrhyparodes and Hileithia and hairlike in all

other genera examined; the uncus is broadly attached

to the tegumen, that is, the tegumen grades evenly into

the uncus without “shoulders”. Character 44:0, shape of

uncus conical, non-capitate, is absent in several species

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

to acute apex; the sacculus is weakly developed or ab-

sent; the bula is long and emerges from near costa base,

pointing towards the centre of the ventral valva edge, or

the bula is reduced to a fold or lost entirely (Crypto-

botys, Pilocrocis, some Herpetogramma spp.); the juxta

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.

The phallus coecum is short or absent; the phallus ap-

odeme is membranous apart from a ventral longitudinal

sclerotized strip; the vesica is granulose, often with a

dense patch of small cornuti. The corpus bursae is mem-

branous or posteriorly with a granulose area, and the

signum is single or absent: when most developed, it is

a round, granulose rhomboid with transverse axis domi-

nant (Cryptobotys, Herpetogramma spp.), or reduced to

a transverse line (Pilocrocis ramentalis Lederer, 1863,

Blepharomastix ranalis), a round dome, or a longitudinal

elongate signum (Hileithia spp.); the signum is absent or

rudimentary in Eurrhyparodes, but with posterior wall of

corpus bursae sclerotized; the corpus bursae is well dis-

tinguished from the long, slender, membranous or partly

sclerotized ductus bursae; the colliculum is membranous

or with a sclerotisation partially encompassing the duc-

tus; the antrum is weakly to strongly sclerotized, simple.

Systematics. Based on our phylogenetic results, we place

Eurrhyparodes Snellen, 1880 (12 spp.), Herpetogramma

Lederer, 1863 (100 spp.) and Hileithia Snellen, 1875 (19

spp.) here. Furthermore, Blepharomastix Lederer, 1863

(85 spp.), Cryptobotys Munroe, 1956 (2 spp.) and Pi-

locrocis Lederer, 1863 (65 spp.) are assigned to Herpe-

togrammatini based on morphological characters.

Munroe (1995) further places the monotypic Pelinop-

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

comprises Acanthaceae (Hileithia, Pilocrocis, Eurrhyp-

arodes splendens Druce, 1895), Actinidiaceae (Pilocro-

cis), Malvaceae (Hileithia) and Urticaceae (Pilocrocis)

(Heppner 2003; solis 2008; Janzen & HallwacHs 2009).

‘Pilocrocis’ milvinalis (Swinhoe, 1886) is reported from

Apocynaceae, Fabaceae and Rubiaceae, P. pterygo-

dia Hampson, 1912 from Lamiaceae (robinson et al.

2010). Eurrhyparodes bracteolalis (Zeller, 1852) is re-

corded from Solanum (Solanaceae) and Oryza (Poaceae)

(robinson et al. 2010), Blepharomastix ranalis from

Chenopodium (Amaranthaceae) (solis 2008). The spe-

cies-rich genus Herpetogramma (100 spp.; nuss et al.

2003 – 2019) contains species with a variety of food

plants ranging from ferns to angiosperms (solis 2008;

Janzen & HallwacHs 2009).

Remarks. The mimetic ‘Pilocrocis’ xanthozonalis

Hampson, 1912 group (including P. cyrisalis (Druce,

1895)) feeds on Rubiaceae; this group is misplaced in Pi-

locrocis and belongs to Agroterini, based on the extended

tegumen, naked uncus, and twin tack-shaped signa.

4.2.13. Hymeniini + Asciodini

Synapomorphies. 61:1, two or more hairpencil sclerites

on each side of the genitalia (articulated with each other

via membranes); 62:1, more than one kind of hairpencil

chaetae present.

4.2.14. Hymeniini Swinhoe, 1900 stat.rev.

Type genus: Hymenia Hübner, 1825

Synapomorphies. 3:1, transverse rim on anterior or me-

sal face of pedicellus in male present; 9:0, size of 3rd

labial palpomere well developed in both sexes; 75:1,

joint of basal valva costa (with vinculum) extended into

an elongate, rod-shaped process. Slow optimization only:

40:1, sclerite present on each pleural membrane of male

segment 8.

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180

The modied antenna base in males is apomorphic:

the pedicellus is erect, long, with oblong scales emerg-

ing from its distal margin (Hymenia perspectalis (Hüb-

ner, 1796)) or medially and posteriorly from its base

(Spoladea recurvalis); the anterior (H. perspectalis) or

medial edge (S. recurvalis) of the pedicellus is raised to

a transverse rim; the basal agellomeres have a pointy

protrusion on anterior side; and the agellum is directed

posteriad, giving the antenna a geniculate appearance.

The parsimony trees do not have 40:1, but they add

61:1, two or more hairpencil sclerites on each side of the

genitalia, articulated with each other via membranes;

62:1, more than one kind of hairpencil chaetae present;

and 99:1, strongly sclerotized colliculum.

Description. The imagines are small (forewing length

about 9 mm) with dark brown wings contrasted with white

forewing markings in the median and postmedian lines,

and a white transverse band in the hindwing; the head

and legs are contrastingly marked. The basal valva costa

is extended into an elongate, ventrad rod that serves as

dorsal joint with the vinculum. The hairpencils are com-

plex, consisting of several sclerotized pads partly with

parallel lines of sclerotized ridges, bearing distinct bun-

dles of long, characteristically bent chaetae; the anterior

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, modied from lewvanicH 1981, Fig. 18. H – I: schematic antennae. — Scale bars: A, G,

H – 5 mm; B – E – 500 µm.

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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.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

182

half of corpus bursae has a short (S. recurvalis) or long

(H. perspectalis) transverse ridged signum, the posterior

half of the corpus bursae granulose (H. perspectalis) or

densely studded with needle-like spikes (S. recurvalis);

the antrum has a longitudinal, non-sclerotized strip.

Hymeniini exhibits a typical wing pattern with a dark

to light brown ground colour interrupted by a broad white

postmedial line on fore- and hindwing; abdomen dorsally

brown with a white band on the segments’ posterior mar-

gin.

Systematics. Based on our phylogenetic analyses, we

place Hymenia Hübner, 1825 (3 spp.) and Spoladea

Guenée, 1854 (2 spp.) in Hymeniini. Hymenia and Spo-

ladea represent a part of Munroe’s (1995) polyphyletic

Hymenia genus group.

Food plants. The spotted beet webworm moth, Hymenia

perspectalis (Hübner, 1796), and the Hawaiian beet web-

worm moth, Spoladea recurvalis, are polyphagous, their

larvae feeding on a large variety of food plants, among

them several important crops like Amaranthus, Beta, So-

lanum tuberosum, Spinacia, Xanthosoma and Zea mays

(solis 2006, 2008).

Remarks. The extension of the basal valva costa into

an elongate, ventrad rod, serving as dorsal joint with the

vinculum, is also present in most investigated Margaro-

niini, in Arthromastix lauralis (Asciodini), Samea ec-

clesialis Guenée, 1854 (Nomophilini), and Prophantis

xanthomeralis (Trichaeini).

The imagines of Hymenia and Spoladea are very sim-

ilar externally, and the generic names have been used in-

terchangeably in the literature. Despite these supercial

similarities between the adults of Spoladea and Hymenia,

their genitalia are signicantly different, and the two gen-

era should be kept separate.

4.2.15. Asciodini Mally, Hayden, Neinhuis,

Jordal & Nuss trib.n.

Type genus: Asciodes Guenée, 1854

Synapomorphies. 5:1, sensillar setae on basal antenno-

meres of male > 50% relative to diameter of basal anten-

nomeres; 72:1, extension (process in some cases) of dor-

sodistal sacculus present; 74:1, basal costa inated; 86:0,

phallus apodeme sclerotisation reduced to a ventral, lon-

gitudinally sclerotized strip; 107:0, corpus bursae with a

granulose sclerotised area. Slow optimization only: 52:1,

connection point of transtillum arms broad; 73:2, bula

and dorsodistal sacculus fused.

The parsimony trees do not have 107:0, but they add

ve synapomorphies: 8:0, dorsal direction of 3rd labial

pal po mere; 33:1, male tergite 8 with heterogenous, dis-

tinct sclerotization pattern; 51:1, transtillum arms rounded;

61:1, two or more hairpencil sclerites on each side of the

genitalia, articulated with each other via membranes; and

62:1, more than one kind of hairpencil chaetae present.

Description. Males of some genera exhibit modied an-

tennomeres halfway along the agellum. In the forewing

of many genera, the postmedial line is roundly concave

where it crosses the anal fold, rather than angulate. The

sacci tympani are hemispherical and clearly dened; they

are smaller and deeper than in most Spilomelinae. The

uncus head is bicapitate or has two separate heads (with a

single head and a central dorsoventral, chaetae-free strip

in Arthromastix lauralis (Walker, 1859) and Ceratocilia

sixolalis (Schaus, 1912)); the costa base and vinculum

saccus are inated; the transtilla is large and circular or

strap-like, with a broad median connection (slim in C. six-

olalis). The bula in the center of the valva is connected

to the sacculus by a distinct “arch” bowing transversely

across the valva. This arch may bear from one to three

digitate processes, or none (Bicilia). Signa are usually ei-

ther absent (most genera) or present as one arcuate line

(Psara, Sathria, Bicilia). The ostium bursae and ductus

bursae are variously sclerotized. Minimally, the collicu-

lum is smooth, elongate, and entire (not ventrally mem-

branous). In some genera, the ostium is anked by two

plates or entirely surrounded by wrinkled sclerites, and

the colliculum may be fused with more extensive scleroti-

zation along the ductus bursae, which is always shorter

than the corpus bursae. The posterior end of the corpus

bursae is often sclerotized with granules or spinules.

Systematics. Based on our phylogenetic results we place

Arthromastix Warren, 1890 (2 ssp.), Asciodes Guenée,

1854 (5 spp.) and Arthromastix pactolalis (Guenée, 1854)

comb.n. here. Furthermore, based on common morpho-

logical features, we assign Beebea Schaus, 1923 (1 sp.),

Bicilia Amsel, 1956 (4 spp.), Ceratocilia Amsel, 1956

(8 spp.), Ceratoclasis Lederer, 1863 (9 spp.), Laniifera

Hampson, 1899 (1 sp.), Laniipriva Munroe, 1976 (1 sp.),

Loxomorpha Amsel, 1956 (4 spp.), Maracayia Amsel,

1956 (2 spp.), Psara Snellen, 1875 (36 spp.) and Sathria

Lederer, 1863 (3 spp.) to Asciodini.

We transfer ve genera from Munroe’s (1995) Siga

group: Beebea, Laniifera, Laniipriva, Loxomorpha and

Maracayia. Males of all the species have the bula con-

nected to the sacculus by an arch, valvae oval in shape

or with a basally inated costa, enlarged transtilla bases

(exept Loxomorpha), and the forewing PM line rounded

basad on the anal fold. Females have a sclerotized la-

mella postvaginalis, except in Laniipriva. The known

larvae feed on Cactaceae (Caryophyllales) as borers or

webworms. The robust form of the genitalia obscures a

key morphological character – the sacculus-bula arch –

but it is visible in careful dissection. Unlike most Ascio-

dini, the hairpencils are either very simple tufts of hairs

or absent, and male antennae are not modied. Like in

Margaroniini and Spilomelini, the large size of imagines

and “robust” genitalia are syndromatic of the internally

feeding larval habit. Laniipriva is problematic because

the female genitalia illustrated by Munroe (1976b: g.

21) have an unarmed ostium and a bulged colliculum like

in the Siga group, but the female maculation (ibid. g. 6)

is typical of Asciodini.

Ceratocilia (considering C. sixolalis) may have a ba-

sal position in this tribe, with its simple transtilla and hair-

pencils.

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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Food plants. Asciodini larvae commonly feed on her-

baceous Caryophyllales. Asciodes gordialis Guenée,

1854 feeds mainly on Nyctaginaceae (Bougainvillea

Mirabilis, Pisonia), as does Ceratocilia sixolalis (Neea,

Pisonia); two undetermined Ceratocilia species were

reared from Rubiaceae, though (kiMball 1965; Janzen

& HallwacHs 2009; robinson et al. 2010). Arthromas-

tix lauralis feeds on Trichostigma octandrum (Phytolac-

caceae) (bendicHo-lopez 1998). Bicilia is recorded from

Petiveria and Rivina (Petiveriaceae) (bendicHo-lopez

1998; Janzen & HallwacHs 2009). Psara feeds on Ama-

ranthaceae, Nyctaginaceae, Nelumbonaceae, Phytolac-

caceae, P. obscuralis also on Convolvulaceae (Janzen

& HallwacHs 2009; robinson et al. 2010). Laniifera

cyclades (Druce, 1895), Beebea guglielmi Schaus, 1923,

Loxomorpha, and Maracayia species feed on Cactaceae,

especially Opuntia, with records of Maracayia on other

Caryophyllales (Mann 1969; Janzen & HallwacHs 2009;

lara-villalón et al. 2016).

Remarks. The upturned palpi (8:0) and heterogenous

male tergite 8 (33:1) are shared with some other taxa (see

diagnoses of Agroterini and Margaroniini). An extension

of the sacculus (72:1) is shared with Eurrhyparodes, but

in that genus, it is a free process, not fused with the bula.

This group of uncolorful moths, as circumscribed

here, is a Neotropical radiation on Caryophyllales. Mun-

roe’s (1995) association of the gracile external feeders is

one of his more perceptive groupings; we doubt that he

had knowledge of the host records available to us now.

Munroe probably associated the large-bodied Beebea and

Laniifera with Siga on overall habitus, and he left Loxo-

morpha and Maracayia unplaced.

Hymeniini species, although polyphagous, prefer

Amaranthaceae and Chenopodiaceae. Therefore, the sis-

ter-group relationship of Asciodini and Hymeniini under

some results (Bayesian and implied-weights parsimony

under k = 9 – 13) suggests that feeding on Caryophyllales

is a synapomorphy of the two tribes.

4.2.16. Trichaeini + (Steniini + Nomophilini)

Synapomorphies. 109:5 (unique), anterior-most signum

a transverse, smooth or dentate arch, with or without cen-

tral posteriad leg (if present, then signum Y-shaped) (not

found with parsimony). Slow optimization only: 33:1,

sclerotization of male tergite 8 heterogeneous (in parsi-

mony trees); 67:2, bula directed towards distal valva;

78:2, general shape of post-basal costa (not the entire

dorsal valva edge) convex (not found with parsimony);

104:0, sclerotisation in ductus bursae absent.

4.2.17. Trichaeini Mally, Hayden, Neinhuis,

Jordal & Nuss trib.n.

Type genus: Trichaea Herrich-Schäffer, 1866

Synapomorphies. 70:1, raised ridge running from basal

to dorsodistal sacculus present.

Description. The valvae are weakly sclerotized, lens-

shaped, often with uting on ventral half of valva (cf.

Odontiinae); the bula is strongly sclerotized (Trichaea)

to weak, bearing simple hairs (absent in some Prophantis

spp.); the sacculus is scaly; a ridge-like protrusion is run-

ning from near the sacculus base to the valva centre. The

corpus bursae has a slim longitudinal signum, its anterior

end split into two anterolateral legs in some Prophantis

species; the ductus bursae is broad, narrowing at the pos-

terior end.

Systematics. Based on our phylogenetic analyses, we

place Prophantis Warren, 1896 (8 spp.) and Trichaea

Herrich-Schäffer, 1866 (11 spp.) in Trichaeini.

Munroe (1967) points to the distinctness of Thlipto-

ceras and Prophantis, with several misplaced species at-

tributable to Prophantis. Our phylogenetic results reveal

that the African T. xanthomeralis is one of these cases.

Here, we remove this and another African species from

Thliptoceras and transfer them to Prophantis: Prophantis

xanthomeralis (Hampson, 1918) comb.n., and Prophant-

is coenostolalis (Hampson, 1899) comb.n. Furthermore,

the African Prophantis longicornalis (Mabille, 1900)

comb.n. is transferred from Syngamia Guenée, 1854.

‘Thliptoceras’ fenestratum Aurivillius, 1910 is also mis-

placed and belongs to one of the non-euspilomeline clad-

es, probably Udeini.

In the parsimony analysis, Desmia falls in Trichaeini,

but it is not supported by any unambiguous morphologi-

cal characters.

A few Neotropical taxa with mimetic maculation and

Rubiaceae-feeding larvae should be investigated as pos-

sible members of Trichaeini, but we leave them incertae

sedis because the morphological evidence is weak and

we did not sequence them. They include Erilusa Walker,

1866 and species misplaced elsewhere, such as Phostria

delilalis (Walker, 1859) and Pilocrocis xanthozonalis

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

relate them to Prophantis. In Erilusa, the uncus is uni-

capitate and the elliptic, dentate bula is not connected to

the sacculus, which exclude it from Asciodini. The uncus

varies from capitate in Erilusa to reduced and triangu-

lar in P. xanthozonalis, but the uncus is likewise variably

reduced in Trichaeini. Sacculosia Amsel, 1956 (1 sp.)

shares a bula and uted valva similar to Trichaea, but

more information is needed in order to investigate this

hypothetical relationship.

Food plants. Prophantis smaragdina (Butler, 1875), P.

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’;

alternative hosts are Tricalysia and Bertiera zaluzania,

Ixora coccinea, Gardenia (Rubiaceae), Duranta plumieri

(Verbenaceae) and Triclisia (Menispermaceae) (waller

et al. 2007; guillerMet 2009). HinckleY (1964) reports

an undescribed Prophantis from Fiji boring in Gardenia

owers and shoots.

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184

Janzen & HallwacHs (2009) report Trichaea larvae

from Psychotria spp., Morinda panamensis and Marga-

ritopsis microdon (Rubiaceae), with a single record on

each Urticaceae and Celastraceae. Feeding on Rubiaceae

is shared with Nomophilini (see also remarks there).

Remarks. The slim longitudinal signum of some Pro-

phantis species with its anterior end split into two ante-

rolateral legs is paralleled in Syngamia (Nomophilini).

4.2.18. Steniini Guenée, 1854 stat.rev.

Type genus: Stenia Guenée, [1845] = Dolicharthria Stephens, 1834

Synapomorphies. 106:0, sclerotisation in corpus bursae

absent. Slow optimization only: 73:0, bula and dorso-

distal sacculus (or its extension) distant from each other,

non-overlapping. The parsimony trees imply the same,

with both characters unambiguous.

Description. Imagines often have long legs; males have

a slender, long abdomen. The uncus is single or bicapitate

(Loxostegopsis, Tatobotys) or entirely split (Metasia) and

has bid chaetae. The valva costa is concave or straight,

in some taxa weakly convex, and the valva is simple with

usually zero or one bula originating from base of valva,

or in the Duponchelia group (sensu HaYden 2011) with

two or three small bulae at the base of the valva; the

phallus has a caecum. The signum is absent, except in

Bradina, Diathrausta, and Perisyntrocha, where it is a

toothed arc. The ostium and ductus bursae lack any other

sclerotization.

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.

185

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In the Duponchelia group, Rs1 is stalked with Rs2+3

in the forewing, and a fovea in the male forewing at the

distal end of the discal cell is a recurrent character in sev-

eral but not all species; a pair of elongate hairpencils is

present dorsal of the vinculum; the ductus bursae is very

short; the larvae have the mesothoracic SD2 seta ne and

hairlike.

Systematics. Based on our phylogenetic results, we

place Anageshna Munroe, 1956 (1 sp.), Dolicharthria

Stephens, 1834 (24 spp.), Duponchelia Zeller, 1847 (5

spp.) and Metasia Guenée, 1854 (88 spp.) in Steniini.

Furthermore, based on morphology we place Apogeshna

Munroe, 1956 (3 spp.), Bradina Lederer, 1863 (87 spp.),

Epherema Snellen, 1892 stat.rev. (1 sp.), Hymenoptychis

Zeller, 1852 (4 spp.), Loxostegopsis Dyar, 1917 (6 spp.),

Penestola Möschler, 1890 (3 spp.), Steniodes Snellen,

1875 (9 spp.), Symmoracma Meyrick, 1894 (1 sp.) and

Tatobotys Butler, 1881 (11 spp.) here. Bradina is unusual

in possessing a signum, which could be plesiomorphic.

The genera Duponchelia, Hymenoptychis, Penestola and

Tatobotys are considered to be closely related: they have

two or three small bulae, a pair of narrow vincular andro-

conia, forewing Rs1 stalked with Rs2+3 and frequently a

fovea, and the larvae are semiaquatic in swamps. We did

not examine the type species of Nacoleia, N. rhoeoalis

(Walker, 1859), but certain important species in this large

genus (84 spp.) such as N. octasema (Meyrick, 1886) and

N. charesalis (Walker, 1859) belong to Steniini based on

the position of the bula, absence of a signum, and sap-

rophagous larval habits.

Piletocera Lederer, 1863 (93 spp.) probably belongs

to Steniini. We have not studied the type species P. viola-

lis Lederer, 1863, but P. signiferalis (Wallengren, 1860)

as illustrated by clarke (1986: gs. 56, 57) shares the

maculation, a broad and deep saccus, ornate valvae, com-

plex hairpencils, and a corpus bursae with spicules but

no single signum. It is related to a group of Steniini that

have a broad saccus and ornate valvae that includes cer-

tain Steniodes species (S. mendica (Hedemann, 1894), S.

acuminalis (Dyar, 1914)), Camptomastix Warren, 1892

and Symmoracma Meyrick, 1894. Lipararchis Meyrick,

1934 (2 spp.) might belong here too.

Food plants. Little is known about the feeding habits of

Steniini. Dolicharthria punctalis (Denis & Schiffermül-

ler, 1775) preferably on wilting leaves of different plants

(Hasenfuss 1960). Metasia corsicalis (Duponchel, 1833)

is reported to feed on detritus (leraut 2012). Nacoleia

charesalis feeds on rotting leaves and bores in turmeric

stems, and N. octasema consumes inorescences of ba-

nanas (paine 1964; HireMatH et al. 1990; kuMar et al.

1996; toMinaga 2002). The absence of records by itself

suggests that the saprophagous habit is common, because

such larvae would be easy to overlook. One group is par-

ticularly interesting: the larvae of the Duponchelia group

(sensu HaYden 2011) are detritivores in marshes and

intertidal environments of mangrove swamps (MurpHY

1990). They are often associated with the Avicennia zone

of mangrove forests, which is inundated at high tides. The

larvae live on the ground and feed on rich soil and juicy

fallen plant matter (e.g. HinckleY 1964). Duponchelia

fovealis Zeller, 1847 as a pest is spread through the plant

nursery trade as it feeds on organic potting soil and suc-

culent stems and foliage.

4.2.19. Nomophilini Kuznetzov & Stekolnikov,

1979 stat.rev.

Type genus: Nomophila Hübner, 1825

Synapomorphies. No unambiguous synapomorphies

could be found for this tribe as circumscribed in the

Bayesian results. Slow optimization only: 74:1, basal

costa inated. Two sister clades are present in Nomophil-

ini, Syngamia + (Ategumia + (Bocchoris + (Diasemia

+ Diasemiopsis))) and Desmia + ((Mecyna + Arnia) +

(Samea + Nomophila)). In the former clade, no synapo-

morphies or characters from slow optimization are found,

but the latter clade is characterised by the synapomor-

phies 67:0, bula ventrally directed towards sacculus or

distal sacculus, and 95:0, longitudinal membranous strip

in the antrum sclerotisation absent.

Nomophilini as circumscribed here is not monophyl-

etic in the parsimony trees.

A core Nomophilini s.str. consisting of Mecyna, No-

mophila, and Samea (without Desmia or Syngamia) has

several synapomorphies in both the Bayesian and parsi-

mony trees: 5:1, sensilla of male antennae elongate; 89:1,

vesica with multiple cornuti; 100:0, colliculum evenly

sclerotized all around, without membranous strip; and

109:1, signum longitudinal and granular.

Description. Small to medium-sized moths. The wing

pattern is reticulated in many Neotropical genera. The

Diasemia group have sacci tympani normally developed

(Bocchoris), small (Ategumia) or absent (Diasemia,

Diasemiopsis), with the fornix tympani in contact with

the tympanic frame all around. The male genitalia have

a conical to capitate uncus (reduced in Ategumia, Dia-

semia and Bocchoris), uni- to bicapitate, uncus head na-

ked or with simple and/or bifurcate chaetae; the valvae

are ovate, mostly with a convex costa; the bula is well-

developed, straight to arched and emerging from near

the costa base (small in Desmia, absent in Bocchoris,

Diasemia and Diasemiopsis). In the female genitalia, the

corpus bursae has a granulose central area or an elon-

gate signum, longitudinal or transverse in orientation,

in Diasemia and Bocchoris invaginated to form a spine;

the colliculum is sclerotized, in Nomophila and ‘Samea’

multiplicalis (Guenée, 1854) with an apomorphic blind

anterolaterad evagination (diverticulum sensu Munroe

1973); the antrum is strongly sclerotized, broad tubular

or barrel-shaped.

Systematics. Based on our phylogenetic results we place

Arnia Guenée, 1849 (1 sp.), Ategumia Amsel, 1956 (10

spp.), Bocchoris Moore, 1885 (31 spp.), Desmia West-

wood, 1832 (89 spp.), Diasemia Hübner, 1825 (13 spp.),

Diasemiopsis Munroe, 1957 (2 spp.), Mecyna Double-

day, 1849 (34 spp.), Nomophila Hübner, 1825 (14 spp.),

Samea Guenée, 1854 (28 spp.) and Syngamia (25 spp.)

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

186

here. Furthermore, we place Crocidocnemis Warren,

1889 (2 spp.), Diacme Warren, 1892 (10 spp.), Diasemi-

odes Munroe, 1957 (4 spp.), Diathrausta Lederer, 1863

(20 spp.), Epipagis Hübner, 1825 (14 spp.), Mimophobe-

tron Munroe, 1950 (1 sp.), Mimorista Warren, 1890 (15

spp.), Niphograpta Warren, 1892 (1 sp.), Nothomastix

Warren, 1890 (5 spp.), Parapilocrocis Munroe, 1967 (3

spp.), Pardomima Warren, 1890 (16 spp.), Perisyntrocha

Meyrick, 1894 (4 spp.), Pessocosma Meyrick, 1884 (4

spp.) and Sameodes Snellen, 1880 (15 spp.) in Nomo-

philini based on morphological characters.

Arnia Guenée, 1849 was synonymized with Stenia

Duponchel, 1845 (a synonym of Dolicharthria Stephens,

1834) by rebel (1901), a decision that was revoked by

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.

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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aMsel (1952). We nd Arnia as sister to Mecyna, and

they share a number of morphological features, such as

shape of uncus and bula, multiple dentiform cornuti in

the phallus, a central granulose area in the corpus bursae,

and a short broad, sclerotized ductus bursae.

Food plants. Larvae are leaf-rollers mainly on Fabaceae,

Onagraceae, Rubiaceae and Vitaceae, with occasional

records from Begoniaceae, Cordiaceae and Malvaceae

(kiMball 1965; allYson 1984; solis 2008; Janzen &

HallwacHs 2009; HaYden 2014). Ategumia feeds mainly

on Melastomataceae, furthermore on Rubiaceae and Big-

noniaceae, with single records on Fabaceae, Piperaceae

and Urticaceae (Janzen & HallwacHs 2009). Boccho-

ris inspersalis (Zeller, 1852) is reported from Malva-

ceae, Fabaceae and Amaranthaceae (gHesquière 1942;

wagner et al. 2008; robinson et al. 2010). Diasemia is

recorded from Asteraceae, Plantaginaceae and Lecythi-

daceae (gHesquière 1942; robinson et al. 2010), Synga-

mia mainly on Rubiaceae, with further records on Acan-

thaceae and Asteraceae (Janzen & HallwacHs 2009),

Pardomima was recorded from coffee (Rubiaceae) (Mar-

tin 1955). Larvae of the monotypic Mimophobetron feed

on different species of Rubiaceae (Janzen & HallwacHs

2009). Nothomastix klossi is recorded from Psychotria

(Rubiaceae) (Miller et al. 2007).

‘Samea’ multiplicalis (Guenée, 1854) and Niphograp-

ta albiguttalis (Warren, 1889), whose larvae are used in

biological control of aquatic weeds, may represent an

aquatic lineage, related to Crocidocnemis, whose larvae

are not known. Furthermore, Diasemiopsis ramburialis

was reported to feed on leaves of the aquatic fern Azolla

liculoides (Salviniaceae) (faraHpour-HagHani et al.

2016).

In the clade Trichaeini + (Nomophilini + Steniini),

the larvae of most Trichaeini and of several Nomophil-

ini (especially the early-diverging Desmia and Synga-

mia) feed on Rubiaceae. Considering this relationship,

we hypothesize that this is the primitive host family for

Nomophilini or maybe for the entire clade Trichaeini +

(Nomophilini + Steniini), and that Nomophilini has radi-

ated onto other hosts. However, larvae of Steniini, as far

as known, are detritivorous.

Remarks. The common form of the male genitalia of

Nomophilini is fairly nondescript, with few striking

characters: the uncus is usually simple or weakly bid,

the valvae are elliptical, and there is one curved bula or

none. However, particular genera or genus groups show

interesting characters, such as modication or loss of the

uncus (Nomophila, Ategumia), distally concave valva

(Sa mea cancellalis, Diacme), or one pair of apical cor-

nuti in several genera (e.g. Mecyna, Mimorista, Samea,

Epi pagis). Likewise, the shape of the signum may be in-

formative above the genus level. The maculation of the

“core” Nomophilini is characteristically chequered, but

other taxa (Desmia, Syngamia, Mimophobetron) show

other patterns. The “core” Nomophilini is well-charac-

terized by female genitalia. The colliculum is tubular

with one or two lateral pockets, and the signa have two

forms: commonly a longitudinal granular strip or (much

less commonly) a transverse “moustache” with two more

or less connected sections, found in Samea castellalis

Guenée, 1854 and Sameodes cancellalis (Zeller, 1852).

An inated basal costa (74:1) is shared with some Ste-

niini (Dolicharthria and Metasia).

Samea is paraphyletic with respect to Nomophila in

our phylogenetic results. Both S. multiplicalis and most

species of Nomophila (see Munroe 1973) have a large,

deeply arched bula, uncus without large bid chaetae,

and the colliculum extended into a diverticulum.

Nomophila was revised by Munroe (1973), the Af-

rican Pardomima species by Martin (1955), Syngamia

orella (Stoll in Cramer & Stoll, 1781) and its variations

by Heppner (2010). This tribe generally corresponds to

the Samea group of Munroe (1995).

A simple tubular colliculum and transverse sig-

num (similar to that in S. castellalis) are also found in

Diasemiodes, Diathrausta, and Perisyntrocha (Munroe

1956), so Nomophilini seems to be a better tribe for these

genera than Steniini. On the other hand, the absence of

checkered maculation suggests that further investigation

is needed.

4.2.20. Munroe’s (1995) Eulepte group

Apart from Syllepte (see below), only the core of Mun-

roe’s (1995) Eulepte group is not represented in our mo-

lecular sampling (three other genera are transferred to

Hydririni). For this reason, we do not formally propose

it as a tribe. The genera Eulepte Hübner, 1825 (6 spp.),

Praeacrospila Amsel, 1956 (4 spp.), Leucochromodes

Amsel, 1956 (8 spp.), and Mesocondyla Lederer, 1863

(2 spp.) have oval to moderately attenuate valvae, uncus

with bid chaetae, and a pair of simple vincular andro-

conia with long, hairlike setae. The saccus is elongate in

Mesocondyla and Eulepte, and perhaps most distinctive-

ly, most taxa (except M. dardusalis) have two inwardly

curved bulae closely set together: one an extension of

the sacculus, the other from the face of the valva, just in-

side and curving in parallel with the saccular bula. The

ductus bursae is elongate in Eulepte and Mesocondyla,

short in Leucochromodes and Praeacrospila, and the sig-

num is absent or double. The maculation is yellow with

a darker postmedial area, which however also occurs

in other taxa (e.g. Lygropia species). Zenamorpha dis-

cophoralis (Hampson, 1899) is another possible member

of this group, considering the male genitalia, although it

could also belong to Trichaeini.

4.2.21. Syllepte Hübner, 1823

Syllepte, the type genus of “Sylleptinae”, is a large poly-

phyletic genus within Spilomelinae, containing 199 valid

species (nuss et al. 2003 – 2019). The identity of the ge-

nus is ambiguous as the type material of its type species,

Syllepte incomptalis Hübner, 1823 (and not Phalaena

amando Cramer, 1779, as erroneously stated by kirti

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

188

& gill 2007), is lost (groll 2017). The illustrations of

the male specimens of this species depicted in Hübner

(1819 – 1823: 18, pl. [50] gs. 285, 286) are difcult to

associate to any known species. The maculation resem-

bles, to some degree, Bocchoropsis Amsel, 1956 and

specimens of the Polygrammodes eleuata (Fabricius,

1777) species group.

4.2.22. Genera removed from Spilomelinae

Aporocosmus Butler, 1886 is transferred to Odontiinae,

where it is related to Thesaurica Turner, 1915. Orthora-

phis Hampson, 1896 is transferred to Lathrotelinae.

Hydropionea Hampson, 1917, Plantegumia Am-

sel, 1956 and Munroe’s (1995) “undescribed genus ex

Boeotarcha Meyrick” appear to form a group of aber-

rant Glaphyriinae. Munroe (1995) probably placed them

in Spilomelinae because the males have the gnathos re-

duced to a transverse band or absent. However, the males

possess a retinacular hook, but they do not have the syna-

pomorphies of Pyraustinae. We transfer them to Glaphy-

riinae s.l. (regier et al. 2012). This is supported by 1)

narrow valvae with apically separate costa and sacculus,

and 2) tympanal organs with large, mesal sacci tympani

and large puteoli. A species of Hydropionea has been

raised on Capparis uniora (Janzen & HallwacHs 2009),

which ts with Glaphyriinae s.l., a clade best dened as

a radiation on mustard-oil producing Brassicales (regier

et al. 2012).

Phaedropsis leialis (Dognin, 1906) and Lygropia

murinalis Schaus, 1912 are related and misplaced in Spi-

lomelinae. They have a male retinacular hook, tympanal

organs with the fornix at the same level as the venula pri-

ma, gnathos with medial process, unmodied valvae, and

an ediacaroid signum. This combination of characters is

very puzzling; we tentatively place them in Pyraustinae

incertae sedis. The host (Gouania Jacq.: Rhamnaceae;

Janzen & HallwacHs 2009) is not informative.

Certain species belong to Pyraustinae incertae se-

dis. Lygropia fusalis Hampson, 1904 and related species

are Pyraustinae, based on the editum of comb-tipped

scales on the sella, concave costa, the deeply invaginated

sacci tympani, and data from the nuclear EF-1a gene.

Blepharomastix haedulalis (Hulst, 1886) is another with

typically pyraustine male genitalia. Females of both taxa

have no signum, so their placement in Pyraustinae was

overlooked.

4.3. Pyraustinae Meyrick, 1890

Type genus: Pyrausta Schrank, 1802

Synapomorphies. 22:1, fornix tympani surface recessed

within the frame (unique); 33:1, heterogenous scleroti-

zation of male tergite 8, i.e. with distinct sclerotization

pattern; 55:1, juxta split 10 – 60% of its length (only in

the parsimony trees); 60:1, partly sclerotized chaetose

hairpencils articulating with the anterior edge of the vin-

culum tegumen connection present; 99:1, strongly scle-

rotized colliculum anterior of the antrum and posterior of

the attachment of the ductus seminalis present (not found

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-

ginate (only in the parsimony trees); 109:6, anterior-most

signum broad, medially constricted, resembling puck-

ered lips (unique; not found with parsimony).

Description. A retinacular hook (frenulum hook sensu

forbes 1926) is present in the male forewing of 13 of the

18 investigated Pyraustinae. The mesothoracic tibia in

males has a hidden hairpencil (oHno 2000; frolov et al.

2007). The fornix tympani is recessed within tympanic

frame. The hemispherical sacci tympani tend to be large

and deep, especially in Pyraustini and Portentomorphini.

The shape and large size is paralleled in some Odontiinae.

The degree at which the praecinctorium of the tympanal

organ is bilobed is neither distinctive for Spilomelinae

nor for Pyraustinae, so that this character is unreliable for

distinguishing the two subfamilies. Male genitalia have

a transtilla inferior (sensu Marion 1954; absent from Te -

tridia and many other taxa) and a sella (sensu Marion

1952) on the inner surface of the valva, often with strong

piliform or spatulate hairs (editum sensu Marion 1952);

the editum is absent from many taxa. The female genita-

lia have a long, coiled ductus bursae (absent in several

taxa, e.g. Nascia, Ostrinia, Uresiphita). Deciduous cor-

nuti are present.

Appendix bursae present, emerging from the anterior

ductus bursae (and not from the corpus bursae) in Te-

tridia, Euclastini and Portentomorphini, or laterally from

the corpus bursae in Pyraustini and Uresiphita. Signum

broad rhombical (Pyraustini), ‘puckered lips’-shaped in

Euclastini and Tetridia Warren, 1890, or ediacaroid in

Portentomorphini and Uresiphita.

The shapes of the teguminal ridges in solis & Maes

(2003: character 9) seem to be good for diagnosing tribes.

Remarks. Plesiomorphic characters shared with the non-

euspilomeline clades in Spilomelinae are: absence of a

sclerotized strip on the pleural membranes of segment 8

(present in Euclastini); costa straight to concave; saccus

of vinculum broadest at the base, without a basal con-

striction. Deciduous cornuti are paralleled in the Spilo-

melinae ‘Syllepte’ adductalis (Walker, 1859) and Pyc-

narmon pantherata (Agroterini). The ediacaroid signum

of Portentomorphini and Uresiphita is shared with the

non-euspilomeline Spilomelinae.

4.3.1. Tetridia Warren, 1890

Autapomorphies. 29:1, large, oval pleural scale tufts

on each side of the male abdominal segment 7 present,

with an opening in its anterior centre (unique); 57:1, ba-

sal saccus constricted; 58:1, ratio between saccus length

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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. vi na cealis.

G: female genitalia of E. splendidalis. H: female genitalia of T. vinacealis. — Scale bars: A, B – 5 mm; C – H – 500 µm.

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

190

and sacculus breadth > 1; 63:1, presence of a pair of

sclerotized, hair-studded hairpencils articulating with

the anteromedian edge of the saccus; 66:0, general shape

of bula broadly triangular; 72:1, extension (process in

some cases) of dorsodistal sacculus present; 76:1, long,

sometimes loosely arranged chaetae on surface of costal

base present; 86:1, phallus apodeme sclerotisation re-

duced to a ventral, longitudinally sclerotized strip; 87:1,

a distinct sclerite in the posterior phallus apodeme pre-

sent; 89:0, vesica with single cornutus; 92:1, ventral end

of papillae anales larger than dorsal end; 100:0, longitu-

dinal membranous strip in the colliculum sclerotisation

absent; 108:1, two or more signa; 110:1, second signum

(located posterior of rst signum) slim, strip-like. Slow

optimization only: 25:0, venulae secundae convergent;

49:1, uncus attached to tegumen as a broad, smooth tran-

sition; 90:2, orientation of everted papillae anales poste-

riad; 112:1, appendix bursae present on anterior ductus

bursae.

Description. The antennae are longer than the forewing.

The uncus has spatulate chaetae in addition to simple,

hair-like chaetae; the sacculus broad and triangular, oc-

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

further towards the distal sacculus; the ventral vinculum

anteromedially has paired hairpencil-like structures. The

signum is broad, medially constricted, resembling puck-

ered lips (‘spectacles-shaped’ in popescu-gorJ & con-

stantinescu 1977); the appendix bursae emerges at the

anterior end of the ductus bursae. The tegumen ridges

cannot be discerned, because the scale-bearing lateral

elds of the tegumen are expanded and compress the me-

sal area into a narrow strip. Therefore, the ridges could be

either absent or fused.

Systematics. So far, we place only two species of Te -

tridia here, the type species T. vinacealis (Moore, 1877),

and T. caletoralis (Walker, 1859).

The identity of T. caletoralis is still not fully clear,

as the type material at NHMUK could not be traced. We

used the DNA extract of voucher specimen WPH209 from

Haines & rubinoff (2012) for our molecular dataset, but

specimens available for morphological study could not

be conrmed as conspecic with specimen WPH209,

which had no genitalia left for investigation. Instead, for

the morphological investigation we used DNA-barcoded

material that was in the nearest neighbour BIN of speci-

men WPH209 in the Barcode of Life Database (BOLD).

kirti & gill (2007) transferred T. caletoralis to Pata-

nia, based on material from the Natural History Museum

London. We have seen the NHMUK Pyralidae slide no.

19900 (male) to which kirti & gill (2007) likely refer

and agree that this taxon belongs to Patania or at least to

Agroterini. However, we have doubts about the correct

identication of the material referred to by kirti & gill

(2007), and we have not seen the specimen from which

NHMUK Pyralidae slide no. 19900 originates. sHibuYa

(1928) mentions four characters in which T. vinacealis,

the type species of Tetridia, differs from T. caletoralis:

body and wings fuscous; legs ferruginous; both wings

with a series of terminal black spots; ante- and postme-

dian lines on the forewing distinctly different in the dor-

sal half. kirti & gill (2007), on the other hand, state that

“this species [T. caletoralis] drastically differs from the

type species of the genus Tetridia Warren i.e., vinacealis

Moore” (kirti & gill 2007: p. 266). Many adult Pyraus-

tinae and Spilomelinae exhibit a wing pattern similar to

that of T. vinacealis, and we assume that kirti & gill

(2007) misidentied their material. A revision of the ge-

nus might bring certainty on this matter and might an-

swer the question whether this taxon should be placed in

a separate tribe.

Food plants. Tetridia caletoralis is recorded from Sho-

rea robusta (Dipterocarpaceae) (robinson et al. 2010).

Remarks. The paired hairpencil-like structures attached

anteromedially to the vinculum are a plesiomorphy

shared with Lamprosema in Spilomelinae: Hydririni; the

‘puckered lips’-shaped signum is shared with Euclastini;

the attachment of the appendix bursae to the anterior end

of the ductus bursae is shared with Euclastini and Porten-

tomorphini.

4.3.2. Euclastini Popescu-Gorj & Constanti-

nescu, 1977 stat.rev.

Type genus: Euclasta Lederer, 1855

Synapomorphies. 10:1, length of maxillary palpi minute

to obsolete, cannot hypothetically come in contact with

each other (not found with parsimony); 16:0, metatibial

proximal inner spur shorter than half of tibial segment

between this and the distal spur pair; 47:2, multifurcate

structure of uncus head chaetae (unique); 65:0, absence

of bula emerging from dorsal valva base near costa base

(not found with parsimony); 78:2, general shape of post-

basal costa (not the entire dorsal valva edge) convex.

Slow optimization only: 18:1, scale brush at costal base

of forewing underside in males formed into a retinacular

hook (not found with parsimony); 44:1, shape of uncus

capitate (not found with parsimony); 49:1, attachment of

uncus to tegumen broad, smooth transition; 109:6, shape

of anterior-most signum broad and medially constricted

(only in the parsimony trees); 112:1, appendix bursae

present on anterior ductus bursae.

Description. The imagines are long-legged and gracile

with narrow, apically rounded forewings with brown

dorsal ground colour, traversed by a whitish band from

wing base to apex, and two dark discal spots. The fore-

wings are held parallel to the frontally raised body when

resting, somewhat resembling Lineodes (Spilomelinae:

Lineodini). The uncus has a bulbous head with multid

chaetae; the valvae are trapezoid, with the straight ven-

tral valva edge parallel to the straight costa that spans the

basal half of the dorsal valva edge; and the distal dorsal

valva edge runs more or less straight towards valva apex.

The bula is absent. The signum is broad, medially con-

stricted and laterally slimly extended, resembling puck-

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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ered lips (‘spectacles-shaped’ in popescu-gorJ & con-

stantinescu 1977); the appendix bursae emerges at the

anterior end of the ductus bursae.

The tegumen ridges are closely parallel, connected by

anterior crossbar at junction of V, like a two-legged “Y”.

Systematics. Euclasta (17 spp.) is the only included ge-

nus.

Food plants. All known host plant records are from Apo-

cynaceae, with Euclasta splendidalis (Herrich-Schäffer,

1848) on Periploca graeca, E. warreni Distant, 1892 on

Acokanthera oppositifolia and E. maceratalis Lederer,

1863 on Gymnanthera nitida (popescu-gorJ & constan-

tinescu 1977; coMMon 1990).

Remarks. The valva shape is somewhat paralleled

in Chilopionea Munroe, 1964 and some species of

Chilochroma Amsel, 1956 (Pyraustini). The ‘puckered

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.

Maes (2000) postulates a close relationship between

Paschiodes Hampson, 1913, Duzulla Amsel, 1952 and

Euclasta. We have not studied Duzulla and the gure and

description in aMsel (1952) are inconclusive, and we

therefore refrain from speculation. We have seen material

of Paschiodes, and the presence of bid chaetae (instead

of multid chaetae as in Euclastini) makes this relation-

ship unlikely. Only one of the ve species of Paschiodes

(P. ugandae Maes, 2005) exhibits an appendix bursae,

and this emerges from the side of the corpus bursae, a

character corresponding to Pyraustini and to Uresiphita

(see remarks under Portentomorphini). Until an analysis

on the phylogenetic relationship of Paschiodes is done,

we refrain from placing the genus in one of the proposed

tribes.

Saucrobotys resembles Euclasta in the bulbous un-

cus head, the valva shape and the absence of a bula and

sella in the male genitalia, and in the appendix bursae

emerging from the anterior end of the ductus bursae as

well as the ‘puckered lips’-shaped signum. The uncus

chaetae are bisetose and not multisetose as in Eucla-

sta. Saucrobotys larvae also feed on Apocynaceae, with

S. futilalis (Lederer, 1863) on Apocynum and Asclepias

syriaca, where the larvae live gregarious in a nest made

from leaves and silk; the plant associations for larvae

and pupae of S. fumoferalis (Hulst, 1886) are doubtful

(Munroe 1976a). Because of the bisetose uncus chaetae,

we refrain from placing Saucrobotys in Euclastini. Eu-

clasta has been revised by popescu-gorJ & constanti-

nescu (1977), the two species of Saucrobotys are treated

in Munroe (1976a).

4.3.3. (Portentomorphini + Uresiphita) +

Pyraustini

Synapomorphies. 20:1, splitting of praecinctorium

weak to absent. 25:1, Course of venulae secundae par-

allel or diverging in posterior half (only in parsimony

trees); 65:1, bula emerging from dorsal valva base near

costa base (only in parsimony trees); Slow optimization

only: 109:3, anterior-most signum transverse rhombical

to cross-shaped, with longitudinal axis shorter than or

equally long as transverse one.

4.3.4. Portentomorphini + Uresiphita

Synapomorphies. 46:0, chaetae on surface of uncus

head(s) absent; 61:1, two or more hairpencil sclerites on

each side of the genitalia (articulated with each other via

membranes); 82:1, costa detached from valval area, the

costa protruding freely dorsad (unique). Slow optimiza-

tion only: 21:1, lobulus on lateral edge of tympanal case

present; 44:1, shape of uncus capitate; 114:1, posterior

point of attachment of appendix bursae on corpus bursae.

4.3.5. Portentomorphini Amsel, 1956 stat.rev.

Type genus: Portentomorpha Amsel, 1956

Synapomorphies. 5:1, sensillar setae of males at basal

antennomeres > 50% relative to diameter of basal anten-

nomeres (not found with parsimony); 46:0, chaetae ab-

sent from surface of uncus head(s); 67:3, bula generally

oriented dorsally towards tegumen or uncus. Slow opti-

mization only: 51:3, transtillum arms large rectangular,

medioventrally with nger-like process (transtilla infe-

rior sensu Marion 1954). The parsimony trees add two

characters: 21:1 and 82:1.

Description. The male genitalia have the costa detached

from the valva and projecting freely dorsad, bearing a

terminal eld of setae. A thin, elongate, curved, often

articulated bula emerges from the centre of the dorsal

valva edge, reaching dorsad; the actual valva consists of

the far dorsad reaching sacculus which ends in a termi-

nal setose eld in the valva apex; the sacculus is large

and membranous. The uncus is narrow, naked, and often

distally forked. In addition to these synapomorphies, the

appendix bursae emerges at the anterior end of the ductus

bursae close to the transition into the corpus bursae, in

Pioneabathra J.C. Shaffer & Munroe, 2007 laterally at-

tached to the corpus bursae. The signum is a four-armed

star in Hyalobathra Meyrick, 1885 and Cryptosara E. L.

Martin in Marion, 1957, an ediacaroid sclerite in Por-

tentomorpha, and in Pioneabathra and Isocentris lalis

(Guenée, 1854) there are two large, opposing granulose

areas. The maculation is basically yellow but often has a

distinctively red or orange postmedial area (or entirely

pink: e.g. Hyalobathra unicolor (Warren, 1895)).The

tegumen is short and evenly sclerotized, without dorsal

ridges.

Systematics. Based on our phylogenetic results we place

Hyalobathra (21 spp.), Cryptosara (3 spp.) and Porten-

tomorpha (1 sp.) in Portentomorphini. Munroe (1976a)

recognizes a group of related genera comprising Porten-

tomorpha, Cryptosara, Isocentris Meyrick, 1887 (7 spp.)

and Hyalobathra. We concur with Munroe’s (1976a)

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

192

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 Anania coronata. G: female genitalia of H. illectalis. — Scale bars: A, B – 5 mm; C – G – 500 µm.

193

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

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composition of this group, here dened as Portentomor-

phini, and further include the African monotypic genus

Pioneabathra.

Food plants. Most food plant records are from Phyllan-

tha ceae (Malpighiales): The monotypic Portento mor pha

feeds on Margaritaria nobilis, Hyalobathra species on

Glo chi dion and Phyllanthus, but they are also record ed

from Abrus (Fabaceae) and Helianthus (Asteraceae); Iso-

centris lalis (Guenée, 1854) and the monotypic Pionea-

bathra on Flueggea, the latter also on Solanum (sutrisno

& Horak 2003; Janzen & HallwacHs 2009; robinson et

al. 2010).

Remarks. The origin of the appendix bursae at the ante-

rior end of the ductus bursae is shared with Tetridia and

Euclastini.

We consistently nd Uresiphita Hübner, 1825 to be

sister to Portentomorphini in our phylogenetic analyses.

Uresiphita does not share any of the unusual synapo-

morphies of the other Portentomorphini, and we there-

fore do not include it in the tribe. Slow optimization in

WinClada results in the following three synapomorphies:

5:1, length of sensillar setae at basal antennomeres rela-

tive to diameter of basal antennomeres (male) > 50%;

21:1, lobules on lateral edge of tympanal case present;

54:2, depth of gap or split of juxta > 60% of dorsoventral

length of juxta to complete division into two juxta arms.

Furthermore, Uresiphita shares with Portentomorphini

the elongate ediacaroid signum. Species of Uresiphita

mainly feed on Fabaceae (Munroe 1976a).

Herpetobotys Maes, 2001 (3 spp.) shares the ediacar-

oid signum and the emergence of the appendix bursae at

the anterior ductus bursae with Uresiphita, but the male

genitalia are different. For now, we leave Uresiphita and

Herpetobotys incertae sedis in Pyraustinae.

The Australian species of Hyalobathra have been re-

vised by sutrisno & Horak (2003).

4.3.6. Pyraustini Meyrick, 1890 stat.rev.

Type genus: Pyrausta Schrank, 1802

= Botydes Blanchard, 1840

= Ennychites Duponchel, 1845

Synapomorphies. 9:0, third labial palpomere well de-

veloped in both sexes; 78:1, general shape of post-basal

costa (not the entire dorsal valva edge) straight; 113:1,

appendix bursae on corpus bursae present. Slow optimi-

zation only: 18:1, retinacular hook present in males. The

parsimony trees imply a very different diagnosis: 5:0,

basal antennomeres of male with sensillae ≤ 50% their

diameter; 10:0, maxillary palpi long enough to contact

each other; 44:0, uncus conical, not capitate.

Description. The uncus is broad to elongate conical,

without a prominent neck constriction and bulbous head;

the uncus has ne setae or robust bifurcate chaetae; many

taxa with a lobate process (sella sensu Marion 1952) on

the central inner valva, carrying long monolament or

multid chaetae (editum sensu Marion 1952). The sig-

num is broad and rhombical, with the transverse axis

longer than the longitudinal axis. The tegumen ridges are

parallel, widely spaced, and not connected.

Systematics. According to our phylogenetic results, we

place the following taxa in Pyraustini: Achyra Guenée,

1849 (19 spp.), Anania (117 spp.), Hyalorista Warren,

1892 (5 spp.), Loxostege (90 spp.), Oenobotys Munroe,

1976 (5 spp.), Ostrinia Hübner, 1825 (21 spp.), Pagyda

Walker, 1859 (26 spp.), Paracorsia Marion, 1959 (1

sp.), Psammotis Hübner, 1825 (8 spp.), Pseudopyrausta

Amsel, 1956 (6 spp.), Pyrausta (341 spp.) and Sitochroa

Hübner, 1825 (10 spp.).

Based on morphology, we furthermore place the fol-

lowing genera in Pyraustini: Adoxobotys Munroe, 1978

(3 spp.), Aglaops Warren, 1892 (4 spp.), Anamalaia

Munroe & Mutuura, 1969 (1 sp.), Arenochroa Munroe,

1976 (1 sp.), Aurorobotys Munroe & Mutuura, 1971

(2 spp.), Callibotys Munroe & Mutuura, 1969 (3 spp.),

Carminibotys Munroe & Mutuura, 1971 (1 sp.), Ceutho-

botys Munroe, 1978 (1 sp.), Chilochroma Amsel, 1956

(4 spp.), Chilocorsia Munroe, 1964 (1 sp.), Chilopionea

Munroe, 1964 (1 sp.), Circobotys Butler, 1879 (19 spp.),

Crocidophora Lederer, 1863 (24 spp.), Crypsiptya Mey-

rick, 1894 (8 spp.), Cybalobotys Maes, 2001 (3 spp.),

Deltobotys Munroe, 1964 (3 spp.), Demobotys Munroe

& Mutuura, 1969 (2 spp.), Ecpyrrhorrhoe Hübner, 1825

(12 spp.), Epicorsia Hübner, 1818 (9 spp.), Epiparbattia

Caradja, 1925 (2 spp.), Eumorphobotys Munroe & Mu-

tuura, 1969 (2 spp.), Fumibotys Munroe, 1976 (1 sp.),

Gynenomis Munroe & Mutuura, 1968 (2 spp.), Hahn-

cappsia Munroe, 1976 (39 spp.), Helvibotys Munroe,

1976 (5 spp.), Limbobotys Munroe & Mutuura, 1970

(5 spp.), Munroeodes Amsel, 1957 (4 spp.), Nascia J.

Curtis, 1835 (3 spp.), Neadeloides Klima, 1939 (2 spp.),

Neoepicorsia Munroe, 1964 (7 spp.), Neohelvibotys

Munroe, 1976 (9 spp.), Nephelobotys Munroe & Mu-

tuura, 1970 (1 sp.), Nomis Motschulsky, 1861 (4 spp.),

Oronomis Munroe & Mutuura, 1968 (1 sp.), Palepicor-

sia Maes, 1995 (1 sp.), Paranomis Munroe & Mutuura,

1968 (4 spp.), Paratalanta Meyrick, 1890 (9 spp.), Par-

battia Moore, 1888 (6 spp.), Perispasta Zeller, 1876 (1

sp.), Placosaris Meyrick, 1897 (20 spp.), Powysia Maes,

2006 (1 sp.), Prooedema Hampson, 1896 (1 sp.), Protepi-

corsia Munroe, 1964 (13 spp.), Pseudepicorsia Munroe,

1964 (4 spp.), Pseudognathobotys Maes, 2001 (2 spp.),

Pseudopagyda Slamka, 2013 (3 spp.), Pseudopolygram-

modes Munroe & Mutuura, 1969 (1 sp.), Pyrasia M. O.

Martin, 1986 (1 sp.), Sarabotys Munroe, 1964 (2 spp.),

Sclerocona Meyrick, 1890 (1 sp.), Sinibotys Munroe &

Mutuura, 1969 (5 spp.), Thivolleo Maes, 2006 (4 spp.),

Thliptoceras Warren, 1890 (31 spp.), Toxobotys Munroe

& Mutuura, 1968 (3 spp.), Vittabotys Munroe & Mutuu-

ra, 1970 (1 sp.) and Xanthostege Munroe, 1976 (2 spp.).

Food plants. Achyra, Anania, Hyalorista, Loxostege

and Sitochroa are polyphagous on a variety of host

plants (Munroe 1976a; robinson et al. 2010; Janzen

& HallwacHs 2009). The Central American species of

Pyrausta mainly feed on Lamiaceae, Verbenaceae, Ama-

ranthaceae (Janzen & HallwacHs 2009). Oenobotys is re-

corded from Eupatorium (Asteraceae) (Munroe 1976a).

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

194

Pagyda species feed on Verbenaceae and Scrophulari-

aceae, Psammotis on Lamiaceae, and Pseudopyrausta

on Lantana (Verbenaceae) (robinson et al. 2010). The

monotypic Paracorsia in mainly found on Fabaceae

(Genista, Ulex, Cytisus, Phaseolus), but also on Scrophu-

lariaceae (Verbascum) (lHoMMe 1935).

Remarks. Our concept of Pyraustini still comprises

the majority of the genera and species of Pyraustinae.

Although we do not further subdivide the tribe, many

characters would provide good evidence. For example,

the spatulate scales of the editum characterize many gen-

era. Achyra, Loxostege, Powysia and Sitochroa share a

unique apomorphic anterior directed projection medially

on the frons (1:1). Species of Anania share a unique syn-

apomorphic cone-shaped central structure in the antrum

(97:1) (leraut 2005a; tränkner et al. 2009).

5. Discussion

The phylogenetic analysis of our dataset results in the

same relationships among Crambidae as found by regier

et al. (2012) based on a different set of molecular markers

that overlaps with our dataset in part of the CAD gene.

Because of the low number of Crambidae outgroup taxa

in our dataset, comparison with the topology of regier et

al. (2012) is only possible to a limited degree, but our re-

sults reect their ndings where the “Wet Habitat Clade”

is sister to the clade of Crambinae + Scopariinae.

regier et al. (2012) did not include a representative

of the Sufetula genus group in their analysis. We include

Sufetula in our dataset, and we nd it to fall outside of

Spilomelinae, agreeing HaYden (2013) and Minet (2015)

who argued for the exclusion of the Sufetula group from

Spilomelinae. Instead, Sufetula is sister to the “CAMMSS

clade” minus Musotiminae sensu regier et al. (2012).

Minet (2015) re-established the name Lathrotelinae on

a subfamily rank for the Sufetula group and placed the

taxon near Acentropinae based on shared characters of

the sternum on abdominal segment A2 and the oviposi-

tor. A phylogenetic analysis of a larger taxon sampling of

Crambidae, including all currently accepted subfamilies,

is necessary to investigate the relationship of Lathroteli-

nae within Crambidae. In his morphology-based phylo-

genetic analysis of Australian Spilomelinae, sutrisno

(2002a) nds Diplopseustis, now in Lathrotelinae (Minet

2015), as subordinate in Spilomelinae, and sister to the

monotypic Aboetheta.

Wurthiinae, with the single genus Niphopyralis (=

Wurthia), was originally described in Arctiidae (roepke

1916). keMner (1923) synonymised the group with Sch-

oenobiinae, where Munroe (1958) retained it, while

lewvanicH (1981) transferred it to Pyraustinae (s.l.). re-

gier et al. (2012) found Niphopyralis to be ingroup of

a strongly supported Spilomelinae, and consequently

synonymized Wurthiinae with Spilomelinae. We conrm

that Niphopyralis belongs in Spilomelinae and assign it

to Wurthiini based on our phylogenetic results. As in the

studies of Mutanen et al. (2010) and regier et al. (2012),

this taxon exhibits a very long terminal branch in our

phylogenetic results (Fig. 1). The RogueNaRok analysis

marked Niphopyralis as rogue taxon, but we decided to

keep it in the dataset as we wanted to investigate its rela-

tionship with other Spilomelinae (see 3.3.). Most of the

observed substitutions in N. chionesis relative to other

investigated taxa are synonymous, i.e. they do not cause

a change in the translated amino acid.

We nd little congruence between our phylogenetic

results and those of the study by sutrisno (2002a) based

on 42 external and genital characters of adult moths of

selected Australian Spilomelinae, partly due to the little

taxon overlap between the two datasets. sutrisno (2002a)

proposed two synapomorphies for Spilomelinae, namely

a strongly bilobed praecinctorium and the absence of a

retinacular hook. We nd the former character to not be

consistent among Spilomelinae; the latter character is in-

deed not found among Spilomelinae, but present in most

Pyraustinae, although it is reduced in many taxa. The

common ndings in both phylogenies are: Isocentris +

Hyalobathra, which we place in Portentomorphini; Hy-

menia + Spoladea, both in Hymeniini; Hymenoptychis +

Tatobotys (misspelled as ‘Tatabotys’ in sutrisno 2002a)

are placed in Steniini. Furthermore, Agrioglypta, Chryso-

thyridia, Didymostoma, Dysallacta, Glyphodes, Synclera

and Talanga form a monophylum in sutrisno (2002a),

and we place all seven genera in Margaroniini. We also

nd the synapomorphies proposed by sutrisno (2002a)

among our synapomorphies for this clade, i.e. a heterog-

enous sclerotization of male tergite 8 (character 33:1),

and two signa (character 108:1).

Our phylogenetic results largely reect those of the

study of Haines & rubinoff (2012) on Omiodes. In their

phylogram (g. 2 therein), nine of our proposed Spilo-

melinae tribes can be identied: Udeini (Udea), Agroter-

ini (Patania, Pleuroptya), Spilomelini (Cnaphalocrocis,

species misplaced in ‘Phostria’), Hymeniini (Spoladea),

Herpetogrammatini (Herpetogramma), Trichaeini (Pro-

phantis), Nomophilini (Nomophila, Sameodes), Steniini

(Bradina, Piletocera), and Margaroniini (‘Omiodes’ ba-

salticalis and its sister clade). Interestingly, Haines &

rubinoff (2012) found a clade comprising the still un-

placed genera Prorodes, Syllepte and Coptobasis as sister

to Bradina + Piletocera.

The GENES- and TIGER-partitioned results differ

in topology, the most fundamental difference being the

placement of Spilomelini, which in the GENES-parti-

tioned analyses is sister to Margaroniini (with PP << 0.9;

see dotted line in Fig. 1). Other differences include the

lack of support (i.e. PP < 0.9) in TIGER-partitioned anal-

yses for the monophyla Midila + Schoenobius, (Psammo-

tis + Pseudopyrausta + Anania), and Udeini + Lineodini.

waHlberg et al. (2005) reported synergistic effects

of combined morphological and molecular data for their

phylogenetic analysis of Papilionoidea, and in their re-

view of studies using these two kinds of data, wortleY

& scotland (2006) nd that most often node resolution

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ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

and support increase with the addition of morphological

data to a genetic dataset. In our results, the phylogram is

nearly fully resolved, and most nodes have high support

even in the dataset only comprising the molecular data

(see posterior probabilities below branches in Fig. 1).

The only signicant exception is within the Margaroniini

clade, where resolution is poor, and observed relation-

ships as well as their support vary across the differently

partitioned analyses. In contrast to previous authors, the

addition of morphological data to our molecular dataset

does not result in an increase in topology resolution or

branch support.

Based on the poor performance of preliminary anal-

yses which included the morphological coding of the

Crambidae outgroup, we decided to omit this part of the

data. The problems mainly concerned the convergence

of the parallel MrBayes runs and resulted in ESS < 100

for several parameters and a somewhat different topol-

ogy, where Pyraustinae is sister to a monophylum of

Spilomelinae and the Crambidae outgroup. Due to the

insufcient ESS, this alternative topology was rejected.

Furthermore, the choice and circumscription of the mor-

phological characters focus on Spilomelinae and Pyraus-

tinae, and for many characters, we are not condent about

drawing homologies with other groups of Crambidae. At

the same time, other crambids exhibit characters that

are not present in Spilomelinae and Pyraustinae, e.g. the

well-developed gnathos of Scopariinae, Crambinae, Sch-

oenobiinae, Glaphyriinae and other groups. It is unlikely

to get meaningful results for the phylogeny of Spilomeli-

nae if morphology is coded, for example, based on the

characters dened by landrY (1995) for Crambini, or by

sutrisno (2002b) for the Australian Glyphodes species

and resembling genera. Consequently, a morphomatrix

that covers characters from all Crambidae taxa and that

is based on a less biased taxon sampling would be neces-

sary to better reect the morphological diversity of the

focus group and to lead to more meaningful phylogenetic

results.

In contrast to the other excluded Crambidae outgroup

taxa, we choose to retain the Lathrotelinae Sufetula in the

nal morphological data matrix in order to investigate its

placement in the phylogeny based on all available data.

Sufetula was recently removed from Spilomelinae (Mi-

net 2015), and we concur with this decision as we nd

the genus to not belong to Spilomelinae in our phyloge-

netic results.

Pyraustinae and Spilomelinae are both strongly sup-

ported monophyletic and sister to each other, as found

by regier et al. (2012), but opposed to solis & Maes

(2003) who found the two groups distantly related. The

difference in the structure of the fornix tympani, recessed

within the tympanic frame in Pyraustinae and projecting

ventrally beyond the tympanic frame in Spilomelinae, is

the most consistent character for distinguishing the two

groups, and underlines the importance of the tympana

for pyraloid systematics. forbes (1926: p. 332) mentions

that the absence or presence of the retinacular hook sep-

arates Pyraustinae (s.l.) into two “mainly if not wholly

natural lines”, i.e. Spilomelinae and Pyraustinae sensu

stricto. We concur with forbes (1926), and furthermore

consider the presence of a retinacular hook the plesio-

morphic character state as it is found in a number of taxa

in the sister group of Spilomelinae + Pyraustinae, e.g. in

Scopariinae (nuss 2005), Crambinae (landrY 1995) and

Schoenobiinae (lewvanicH 1981); see sauter (1973) for

a detailed study on this character among Pyraloidea. The

retinacular hook is absent in all investigated Spilomeli-

nae, and it may therefore serve as a diagnostic character

for the group. In other subfamilies, however, the pres-

ence or absence of this structure in males is highly vari-

able at the generic (solis & Maes 2003) or species level

(nuss 2005).

Character 115 (locality of larval feeding) is the only

character of the immature life stage, and the only char-

acter not concerning morphology. We chose to include

this character as we considered it as potentially carry-

ing phylogenetic information. Although data coverage

for this character is only about 40%, some statements

can be made from the data: For most taxa, there is no

apparent association between phylogenetic lineage and

larval feeding locality. The majority of coded taxa has

larvae that feed concealed in rolled or spun leaves or in

a web (character state 115:0). This is the main feeding

locality for Margaroniini larvae, although some (Agath-

odes, Liopasia, Maruca, Terastia) are partly or entirely

borers in stems, branches, owers, pods and/or fruits. An

interesting association of potential phylogenetic value is

the feeding of Dolicharthria larvae on decaying or dead

plant matter (115:5), a behaviour that is observed in other

Steniini as well (see ‘Food plants’ in 4.2.18.).

Larval host plants and feeding modes (internal vs.

external; leaf rolling, leaf webbing etc.) are considered

a useful source for future research on phylogenetic rela-

tionships among Spilomelinae and Pyraloidea in general.

A study by segar et al. (2017) on phylogenetic predic-

tions of host plant use in Pyraloidea and Geometridae

found that host plant preference is phylogenetically rela-

tively conserved in snout moths. The host plants associa-

tions of the taxa studied here support this observation,

e.g. Lineodini almost exclusively feeding on Solanaceae,

Asciodini commonly on Caryophyllales, and most Por-

tentomorphini on Phyllanthaceae.

The majority of investigated Pyraustinae genera is

placed in Pyraustini, a rather homogenous group which

mainly varies in the shape of the valva and of the sclero-

tised processes on the inner valva surface, as well as in

the uncus shape, although the spectrum of uncus varia-

tion is far narrower than in Spilomelinae. Munroe (1995)

was uncertain about the inclusion of the Neotropical

Portentomorpha and the related Old World Hyalobathra

into his concept of Pyraustinae (Pyraustini sensu Munroe

1995). In our phylogenetic results, both genera are part

of Pyraustinae, and they are placed in Portentomorphini.

However, we nd Hyalobathra to be closer related to the

African Cryptosara. Tetridia, the sister to all other phylo-

genetically investigated Pyraustinae, requires taxonomi-

cal revision (see Systematics under 4.3.1.).

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

196

Spilomelinae is found highly supported monophyl-

etic in our analysis, supporting Munroe’s (1995) opin-

ion of Spilomelinae being “at least in large part” mono-

phyletic, while contradicting Minet (1982) and solis &

Maes (2003) who perceived Spilomelinae as a para- or

polyphyletic assemblage. The only uniquely derived apo-

morphy common to all investigated Spilomelinae is the

ventrad projecting fornix tympani, a character present in

Minet’s (1982) circumscription of the group. A (strong-

ly) bilobed praecinctorium, considered diagnostic by Mi-

net (1982) and sutrisno (2002a), is found to be homo-

plastic. It is absent in several investigated Spilomelinae

and present in a number of Pyraustinae, and therefore of

no diagnostic use. According to solis & Maes (2003) a

bilobed praecinctorium is also present in Midilinae. Ab-

sence characters like the lack of chaetosemata, the sub-

costal retinacular hook, and a well-developed gnathos in

males, as well as the large rhombical signum in females

are not exclusive for Spilomelinae. The loss of the reti-

nacular hook is a synapomorphy of Spilomelinae, but it is

paralleled in some Pyraustinae. The character of distinct-

ly tapered spinulae (Minet 1982) was not investigated.

According to allYson (1981, 1984), there are no diag-

nostic morphological characters distinguishing larvae of

Pyraustinae from those of Spilomelinae.

In our phylogenetic results, the tribe Margaroniini

comprises the most sampled species. This could be due

to a sampling bias, but the large number of taxa attributed

to the tribe (currently 67 genera with 1,044 species) sup-

ports this view of Margaroniini being the most diverse

clade in Spilomelinae. Margaroniini is predominantly

tropical and subtropical in distribution. Apart from com-

prising a large number of genera, the tribe also contains

many species-rich genera, like Palpita, Glyphodes, Omi-

odes and Diaphania.

We nd several well-supported relationships among

Margaroniini: the larvae of Liopasia, Terastia and Ag-

athodes all feed on Erythrina, and imagines of the three

moth genera share a similar wing pattern (sourakov et

al. 2015). We nd the three genera to be closely related,

with Liopasia being sister to Agathodes + Terastia, as re-

ported by sourakov et al. (2015). The Glyphodes group,

as circumscribed by sutrisno (2002b) and sutrisno et

al. (2006), further includes Obtusipalpis as well as the

Dichocrocis zebralis species complex. Conogethes pan-

damalis, related to the species complex of the yellow

peach moth C. punctiferalis, is found to be sister to the

Neotropical genus Azochis. They share a similar anatomy

of the uncus, tegumen and valvae, and a similar structure

of the hairpencil scales. The phylogenetic relationships

to species with a highly similar wing pattern, like the

African Marwitzia species (Maes 1998b) or species of

the Neotropical Polygrammodes eleuata complex, is not

known.

The sister group of Omiodes, as investigated by

Haines & rubinoff (2012), is still not known due to the

extensive polytomy in Margaroniini. We can, however,

rule out Cnaphalocrocis as sister group, which we place

in Spilomelini.

The poorly resolved relationships among Margaronii-

ni should be addressed through the choice of better-suited

genetic markers, e.g. DDC which has a substitution rate

similar to COI (waHlberg & wHeat 2008). Furthermore,

the morphological dataset can be improved to incorpo-

rate additional characters, e.g. structure of the hairpencil

pads and scales, shape and sclerotisation patterns of the

valvae, and shape and structure of the tegumen and un-

cus.

With the morphological dataset at hand and the phy-

logenetic results, we start to gain a better understanding

of functional morphology of the genitalia and character

correlations. In most investigated Spilomelinae, the pres-

ence of sclerotised membraneous strips on the pleural

membranes of the male abdominal segment 8 corresponds

with the presence of hairpencils, and we assume that the

strips might serve as muscular attachment sites for retrac-

tion of the hairpencils. In Pyraustinae, this correlation is

absent, and we nd sclerites on the pleurites of the male

segment 8 only in Euclasta, although most pyraustines

exhibit ‘simple’ hairpencils, i.e. a single small hairpencil

pad on each side of the anterior vinculum-tegumen con-

nection, studded with one type of hairpencil scales. The

longitudinal membranous strip in the female genitalia’s

antrum possibly functions as a stretching zone during

copulation, when the male transfers the spermatophore.

This character is present in species with narrow antrum

while it is rarely found in species with a broad antrum;

it occurs in both Spilomelinae and Pyraustinae. We nd

the bula and the distal sacculus often in close spatial as-

sociation, and the ventral valva margin is often less scle-

rotised in this area, suggesting that this complex might

function as a bendable joint during copulation, when the

male clasps the female genital with its valvae. Minet et

al. (2014) identied the same weak exure on the saccu-

lus as a character of Noctuidae s. str. including Dyopsi-

nae. In the future, emphasis should be put on studying

the muscular attachment regions of this supposed point

for valval bending, as well as of the vinculum and tegu-

men (see kuznetzov & stekolnikov 1979a,b). Like for

most other characters studied, investigation of muscula-

ture and nervature is essential to better understand their

function and homology. Furthermore, the ontogenetic

origins of the bula should be investigated as it is not

clear whether the sclerotised protrusions on the inner side

of the valva that are found in most species of Spilomeli-

nae are homologous. We assume that protrusions emerg-

ing from the costa are not homologous with those arising

from the central inner valva, and we consequently code

them as separate characters.

The morphological circumscription of the observed

tribes in Spilomelinae and Pyraustinae allows for the as-

signment of additional taxa in those tribes through mor-

phological investigation, without the strict requirement

of molecular data. These morphological diagnoses allow

the assignment of additional 125 genera to Spilomelinae

tribes, and additional 56 genera to Pyraustinae tribes. 135

genera of Spilomelinae and 103 genera of Pyraustinae re-

main unassigned to any of the proposed tribes. Among the

197

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

unassigned Spilomelinae genera, the most species-rich

are Syllepte (198 spp.), Nacoleia (84 spp.), Pycnarmon

(59 spp.), Dichocrocis (53 spp.) and Mimudea Warren,

1892 (42 spp.), while 76 genera contain only one to two

species, respectively. In Pyraustinae, the most species-

rich unassigned genera are Semniomima Warren, 1892

(15 spp.), Calamochrous Lederer, 1863 (13 spp.) and

Paliga Moore, 1886 (12 spp.). In the European fauna of

Spilomelinae and Pyraustinae, only Uresiphita remains

unplaced as sister to Portentomorphini. In the Nearctic

region, Daulia, Deuterophysa, Eulepte, Microphysetica,

Stenochora and Syllepte are still unplaced. Furthermore,

the majority of genera from Munroe’s (1995) Neotropi-

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

circumscription of the observed tribes in Spilomelinae

and Pyraustinae allows for the assignment of additional

taxa in those tribes through morphological investigation,

without the strict requirement of molecular data. This

should be applied to the 132 Spilomelinae genera and

103 Pyraustinae genera which are not yet assigned to any

of the tribes. A concerted effort among systematists to

morphologically investigate those unplaced genera and

to assign them to the proposed tribes is feasible and de-

sirable. This effort will likely result in the recognition of

taxa that do not t into this system of tribes. Such taxa

can be morphologically coded as well as sequenced for

the six genetic markers used in this study. Their phyloge-

netic placement can then be inferred through a combina-

tion with the data presented here. We therefore provide a

‘modular’ dataset where taxa of interest can be added in

order to rene the circumscription of the proposed tribes

and to widen our understanding of the phylogenetic rela-

tionships of Spilomelinae and Pyraustinae. The resulting

improved understanding of Spilomelinae and Pyrausti-

nae genera is expected to promote taxonomic revisions

of genera and species groups as well as ecological and

applied research on the pyraloids.

6. Acknowledgements

We are grateful to the following colleagues for providing material

and data for our study: Leif Aarvik (Natural History Museum Oslo,

Norway), Stoyan Beshkov (National Museum of Natural History

Soa, Bulgaria), Jason Dombroskie (Cornell University Ithaca, New

York, U.S.A.), Markus Franzén (Helmholtz Centre for Environmen-

tal Research Leipzig, Germany), Will Haines (University of Hawaii

at Manoa, Hawaii, USA), John B. Heppner, Jacqueline Y. Miller

and Deborah Matthews (all McGuire Center for Lepidoptera &

Biodiversity, Gainesville, Florida, USA), Marianne Horak (CSIRO

Australia), Timm Karisch (Museum of Natural History Dessau,

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

M‘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 (Hydri rini) and

Udea (Udeini)

Hymenia group Ercta: Udeini; Spilomela: Spilomelini; Blepharo-

mastix: Herpetogrammatini; Hymenia, Spoladea:

Hymeniini; Anageshna, Apogeshna, Duponchelia,

Loxostegopsis, Parastenia (= Dolicharthria), Penestola,

Steniodes: Steniini; Desmia, Diasemiodes, Diasemiop-

sis, Diathrausta: Nomophilini

Sacculosia

Diaphania group Margaroniini Chromodes

Polygrammodes group Margaroniini

Siga group Cirrhocephalina, Siga, Zeuzerobotys: Spilomelini;

Beebea, Laniifera, Laniipriva: Asciodini

Eulepte group Gonocausta, Ommatospila, Syllepis: Hydri rini Eulepte, Praeacrospila, Leucochromodes,

Mesocondyla

Samea group Nomophilini Stenorista

Psara group Asciodini

Conchylodes group Conchylodes: Udeini Pycnarmon

Syngamia group Marasmia, Salbia: Spilomelini;

Syngamia: Nomophilini

Sufetula to Lathrotelinae (M

2015)

Diaphantania group Wurthiini

unplaced genera Nonazochis (= Conchylodes), Sisyracera, Tanaophysa:

Udeini; Bocchoropsis, Coenostolopsis, Cyclocena

(H  D 2014: syn. of Microthyris), Gypo des:

Agroterini; Analyta, Caprinia, Meroctena, Tyspa nodes:

Margaroniini; Palpusia, Rhectocraspeda: Spilomelini;

Eurrhyparodes: Herpetogrammatini; Bicilia, Loxomor-

pha, Maracayia: Asciodini; Trichaea: Trichaeini; Bra-

dina: Steniini; Bocchoris, Parapilocrocis: Nomophilini

Agrammia, Carthade, Coelorhyncidia, Co-

remata, Daulia, Deuterophysa, Dichocrocis,

Goniorhynchus, Heterudea, Hyalea, Ischnurg-

es, Luma, Mabra, Massepha, Metoeca,

Metraeopsis, Microphysetica, Mimudea,

Pectinobotys, Piletocera, Piletosoma, Platyg-

raphis, Plectrona, Syntrita, Tanao physopsis,

Trithyris

Ennomosia to Glaphyriinae (H

 D 2014); Hydropionea and

Plantegumia to Glaphyriinae;

Orthoraphis to Lathrotelinae

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

198

Germany), Bernard Landry (Natural History Museum Geneva,

Switzerland), Knud Larsen (Dyssegård, Denmark), Norbert Pöll

(Biologiezentrum Linz, Austria), John Rawlins (Carnegie Museum

of Natural History, Pittsburgh, USA), Rodolphe Rougerie (Natural

History Museum Paris, France), Andreas Segerer (Zoological State

Collections Munich, Germany), Stephen Sutton and Charles Vairap-

pan (University of Sabah, Kota Kinabalu, Malaysia), Marja van der

Straten (Nederlandse Voedsel- en Warenautoriteit Wageningen, The

Netherlands), Boyan Zlatkov (Soa University “St. Kliment Ohrid-

ski” Soa, Bulgaria) and Andreas Zwick (Australian National Insect

Collection, CSIRO, Canberra, Australia), and especially the late

Linwood C. Dow (Largo, Florida), whose efforts to collect, curate,

and professionally identify Neotropical pyraloids was especially

useful for making genera available to study through his bequest.

We are thankful to Miss Chung (Sabah Biodiversity Centre,

Malaysia), Prof. Charles S. Vairappan and Stephen Sutton (both

University Malaysia Sabah, Malaysia), and the staff of Sabah Parks

and the University Malaysia Sabah for collecting preparations

and support in Mount Kinabalu National Park. We further thank

Manuela Bartel, Anke Müller and Anja Rauh (all Senckenberg

Natural History Collections Dresden, Germany) as well as Louise

Lindblom (University of Bergen, Norway) for their support in the

DNA labs. We thank Franziska Bauer, Théo Léger, Klaus-Dieter

Klass (all at Senckenberg Natural History Collections Dresden,

Germany) and Simon Segar (University of South Bohemia, České

Budějovice, Czech Republic) for helpful information and com-

ments, Anastasia Garbar (Bergen, Norway) for the translation of

Russian literature, and Koen V.N. Maes (Wetteren, Belgium) for

support with literature.

We thank Charles A. Boring and Paul Skelley (FDACS-DPI)

for reviewing the manuscript for FDACS, and the reviewers M.

Alma Solis and Brian Wiegmann for their valuable comments on

the submitted manuscript.

JEH generously thanks the Rea Postdoctoral Fellowship of the

Carnegie Museum of Natural History, during which he rst inves-

tigated Munroe’s genus groups and dissected a broad study set of

specimens.

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8. Appendix

Checklist of Spilomelinae and Pyraustinae tribes, associ-

ated genera and species numbers:

SPILOMELINAE

Hydririni: Choristostigma Warren, 1892 (10 spp.) — Gonocaus-

ta Lederer, 1863 (4 spp.) — Hydriris Meyrick, 1885 (7 spp.) —

Lamprosema Hübner, 1823 (72 spp.) [polyphyletic] — Nehydriris

Munroe, 1974 (1 sp.) — Ommatospila Lederer, 1963 (3 spp.) —

Rhectothyris Warren, 1890 (1 sp.) — Syllepis Poey, 1832 (7 spp.)

Lineodini: Atomopteryx Walsingham, 1891 (10 spp.) — Euleuci-

nodes Capps, 1948 (1 sp.) — Leucinodes Guenée, 1854 (20 spp.)

[MALLY et al. 2015; misplaced spp. in Asia and Australia] — Li-

neodes Guenée, 1854 (39 spp.) — Neoleucinodes Capps, 1948

(9 spp.) — Proleucinodes Capps, 1948 (4 spp.) — Rhectosemia

Lederer, 1863 (12 spp.)

Udeini: Cheverella B. Landry, 2011 (1 sp.) — Conchylodes

Guenée, 1854 (21 spp.) [paraphyletic?] — Deana Butler, 1879 (1

sp.) — Ercta Walker, 1859 (7 spp.) — Mnesictena Meyrick, 1884

(7 spp.) — Sisyracera Möschler, 1890 (3 spp.) — Tanaophysa War-

ren, 1892 (2 spp.) — Udea Guenée, 1845 (in Duponchel) (214 spp.)

[e.g. MUNROE 1966, INOUE et al. 2008, MALLY & NUSS 2011]

— Udeoides Maes, 2006 (5 spp.)

Wurthiini: Apilocrocis Amsel, 1956 (11 spp.) — Aristebulea

Munroe & Mutuura, 1968 (2 spp.) — Diaphantania Möschler,

1890 (3 spp.) — Mimetebulea Munroe & Mutuura, 1968 (1 sp.)

— Niphopyralis Hampson, 1893 (8 spp.) — Pseudebulea Butler,

1881 (4 spp.)

Agroterini: Aetholix Lederer, 1863 (4 spp.) — Agrotera Schrank,

1802 (24 spp.) [CHEN et al. 2017] — Aiyura Munroe, 1974 (2

203

ARTHROPOD SYSTEMATICS & PHYLOGENY — 77

(1) 2019

spp.) — Bocchoropsis Amsel, 1956 (2 spp.) — Chalcidoptera

Butler, 1887 (15 spp.) — Chilochromopsis Munroe, 1964 (1 sp.)

— Coenostolopsis Munroe, 1960 (3 spp.) — Diastictis Hübner,

1818 (12 spp.) — Framinghamia Strand, 1920 (1 sp.) — Glau-

cobotys Maes, 2008 (1 sp.) — Goliathodes Munroe, 1974 (1 sp.)

— Gypodes Munroe, 1976 (1 sp.) — Haritalodes Warren, 1890 (11

spp.) — Lygropia Lederer, 1863 (68 spp.) — Lypotigris Hübner,

1825 (1 sp.) — Micromartinia Amsel, 1957 (1 sp.) — Microthyris

Lederer, 1863 (7 spp.) — Nagiella Munroe, 1976 (4 spp.) — Neo-

analthes Yamanaka & Kirpichnikova, 1993 (8 spp.) — Nosophora

Lederer, 1863 (26 spp.) — Notarcha Meyrick, 1884 (18 spp.) —

Pantographa Lederer, 1863 (9 spp.) — Patania Moore, 1888 (40

spp.) — Phaedropsis Warren, 1890 (24 spp.) — Phostria Hübner,

1819 (87 spp.) — Phryganodes Guenée, 1854 (26 spp.) — Tetra-

cona Meyrick, 1884 (2 spp.) [CHEN et al. 2017] — Ulopeza Zeller,

1852 (16 spp.)

Margaroniini: Agathodes Guenée, 1854 (16 spp.) — Agrioglypta

Meyrick, 1932 (11 spp.) [SUTRISNO 2002a,b, 2005, SUTRISNO

et al. 2006] — Alytana J. C. Shaffer, & Munroe, 2007 (2 spp.) —

Analyta Lederer, 1863 (10 spp.) — Anarmodia Lederer, 1863 (24

spp.) — Antigastra Lederer, 1863 (2 spp.) — Aphytoceros Mey-

rick, 1884 (3 spp.) — Arthroschista Hampson, 1893 (2 spp.) —

Asturodes Amsel, 1956 (1 sp.) — Azochis Walker, 1859 (16 spp.)

— Botyodes Guenée, 1854 (10 spp.) — Cadarena Moore, 1886 (1

sp.) — Caprinia Walker, 1859 (11 spp.) — Chabulina J. C. Shaffer,

& Munroe, 2007 (2 spp.) — Charitoprepes Warren, 1896 (2 spp.)

— Chrysophyllis Meyrick, 1934 (1 sp.) — Chrysothyridia Munroe,

1967 (2 spp.) — Cirrhochrista Lederer, 1863 (38 spp.) — Colo-

mychus Munroe, 1956 (2 spp.) — Compacta Amsel, 1956 (4 spp.)

— Condylorrhiza Lederer, 1863 (4 spp.) — Conogethes Meyrick,

1884 (16 spp.) [e.g. INOUE & YAMANAKA 2006, SHASHANK

et al. 2015, 2018] — Cydalima Lederer, 1863 (9 spp.) [STRELT-

ZOV 2008, MALLY & NUSS 2010] — Diaphania Hübner, 1818

(95 spp.) [CLAVIJO ALBERTOS 1990] — Didymostoma Warren,

1892 (2 spp.) [SUTRISNO 2002a] spp.) — Dysallacta Lederer,

1863 (3 spp.) [SUTRISNO 2002a] — Endocrossis Meyrick, 1889

(4 spp.) — Eusabena Snellen, 1901 (4 spp.) — Filodes Guenée,

1854 (16 spp.) — Ghesquierellana Berger, 1955 (5 spp.) —

Glyphodella J. C. Shaffer & Munroe, 2007 (3 spp.) — Glyphodes

Guenée, 1854 (156 spp.) [SUTRISNO 2002a,b, 2003, 2006,

SUTRISNO et al. 2006] — Hedyleptopsis Munroe, 1960 (1 sp.)

— Heterocnephes Lederer, 1863 (4 spp.) — Hodebertia Leraut,

2003 (1 sp.) — Hoterodes Guenée, 1854 (5 spp.) — Leucochroma

Guenée, 1854 (6 spp.) — Liopasia Möschler, 1882 (15 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,

1863 (4 spp.) — Nolckenia Snellen, 1875 (1 spp.) — Obtusipalpis

Hampson, 1896 (6 spp.) — Omiodes Guenée, 1854 (98 spp.) [poly-

phyletic, HAINES & RUBINOFF 2012] — Omphisa Moore, 1886

(10 spp.) — Pachynoa Lederer, 1863 (12 spp.) — Palpita Hübner,

1808 (162 spp.) [INOUE 1996, 1997, 1999] — Parotis Hübner,

1831 (37 spp.) — Poliobotys J. C. Shaffer & Munroe, 2007 (1

sp.) — Polygrammodes Guenée, 1854 (78 spp.) — Polygrammop-

sis Munroe, 1960 (1 sp.) — Prenesta Snellen, 1875 (18 spp.) —

Pygospila Guenée, 1854 (10 spp.) — Radessa Munroe, 1977 (2

spp.) — Rhagoba Moore, 1888 (2 spp.) — Rhimphalea Lederer,

1863 (12 spp.) — Sinomphisa Munroe, 1958 (3 spp.) — Sparagmia

Guenée, 1854 (1 sp.) — Stemorrhages Lederer, 1863 (8 spp.) —

Synclera Lederer, 1863 (13 spp.) — Syngamilyta Strand, 1920 (5

spp.) — Talanga Moore, 1885 (9 spp.) [SUTRISNO 2002a,b, 2005,

SUTRISNO et al. 2006] — Terastia Guenée, 1854 (7 spp.) — Tes -

sema J. F. G. Clarke, 1986 (1 sp.) — Tyspanodes Warren, 1891

(20 spp.) — Uncobotyodes Kirti & Rose, 1990 (1 sp.) — Zebronia

Hübner, 1821 (6 spp.)

Spilomelini: Aethaloessa Lederer, 1863 (3 spp.) — Cirrhocephali-

na Munroe, 1995 (5 spp.) — Cnaphalocrocis Lederer, 1863 (27

spp.) — Eporidia Walker, 1859 (1 sp.) — Geshna Dyar, 1906 (1

sp.) — Marasmia Lederer, 1863 (9 spp.) — Marasmianympha

Munroe, 1991 (1 sp.) — Orphanostigma Warren, 1890 (6 spp.)

— Palpusia Amsel, 1956 (10 spp.) — Rhectocraspeda Warren,

1892 (2 spp.) — Salbia Guenée, 1854 (35 spp.) — Scaptesylodes

Munroe, 1976 (2 spp.) — Siga Hübner, 1820 (2 spp.) — Spilomela

Guenée, 1854 (8 spp.) — Zeuzerobotys Munroe, 1963 (1 sp.)

Herpetogrammatini: Blepharomastix Lederer, 1863 (85 spp.)

— Cryptobotys Munroe, 1956 (2 spp.) — Eurrhyparodes Snel-

len, 1880 (12 spp.) — Herpetogramma Lederer, 1863 (100 spp.)

— Hileithia Snellen, 1875 (19 spp.) — Pilocrocis Lederer, 1863

(65 spp.)

Hymeniini: Hymenia Hübner, 1825 (3 spp.) — Spoladea Guenée,

1854 (2 spp.)

Asciodini: Arthromastix Warren, 1890 (2 spp.) — Asciodes

Guenée, 1854 (5 spp.) — Beebea Schaus, 1923 (1 sp.) — Bicilia

Amsel, 1956 (4 spp.) — Ceratocilia Amsel, 1956 (8 spp.) — Cera-

toclasis Lederer, 1863 (9 spp.) — Erilusa Walker, 1866 (3 spp.)

[tentative placement] — Laniifera Hampson, 1899 (1 sp.) — Lani-

ipriva Munroe, 1976 (1 sp.) — Loxomorpha Amsel, 1956 (4 spp.)

— Maracayia Amsel, 1956 (2 spp.) — Psara Snellen, 1875 (36

spp.) — Sathria Lederer, 1863 (3 spp.)

Trichaeini: Prophantis Warren, 1896 (8 spp.) — Sacculosia Amsel,

1956 (1 sp.) — Trichaea Herrich-Schäffer, 1866 (11 spp.) — Ze-

namorpha Amsel, 1956 (2 spp.)

Steniini: Anageshna Munroe, 1956 (1 sp.) — Apogeshna Munroe,

1956 (3 spp.) — Bradina Lederer, 1863 (87 spp.) — Dolicharthria

Stephens, 1834 (24 spp.) — Duponchelia Zeller, 1847 (5 spp.) —

Epherema Snellen, 1892 (1 sp.) — Hymenoptychis Zeller, 1852 (4

spp.) — Loxostegopsis Dyar, 1917 (6 spp.) — Metasia Guenée,

1854 (88 spp.) — Penestola Möschler, 1890 (3 spp.) — Steniodes

Snellen, 1875 (9 spp.) — Symmoracma Meyrick, 1894 (1 sp.) —

Tatobotys Butler, 1881 (11 spp.)

Nomophilini: Arnia Guenée, 1849 (1 sp.) — Ategumia Amsel,

1956 (10 spp.) — Bocchoris Moore, 1885 (31 spp.) — Crocidoc-

nemis Warren, 1889 (2 spp.) — Desmia Westwood, 1832 (89 spp.)

— Diacme Warren, 1892 (10 spp.) — Diasemia Hübner, 1825

(13 spp.) — Diasemiodes Munroe, 1957 (4 spp.) — Diasemiopsis

Munroe, 1957 (2 spp.) — Diathrausta Lederer, 1863 (20 spp.) —

Epipagis Hübner, 1825 (14 spp.) — Mecyna Doubleday, 1849 (34

spp.) — Mimophobetron Munroe, 1950 (1 sp.) — Mimorista Wa r-

ren, 1890 (15 spp.) — Niphograpta Warren, 1892 (1 sp.) — No-

mophila Hübner, 1825 (14 spp.) [MUNROE 1973] — Nothomastix

Warren, 1890 (5 spp.) — Parapilocrocis Munroe, 1967 (3 spp.)

— Pardomima Warren, 1890 (16 spp.) — Perisyntrocha Meyrick,

1894 (4 spp.) — Pessocosma Meyrick, 1884 (4 spp.) — Samea

Guenée, 1854 (28 spp.) — Sameodes Snellen, 1880 (15 spp.) —

Syngamia Guenée, 1854 (25 spp.)

PYRAUSTINAE

Euclastini: Euclasta Lederer, 1855 (17 spp.) [POPESCU-GORJ &

CONSTANTINESCU 1977]

Portentomorphini: Cryptosara E. L. Martin in Marion, 1957 (3

spp.) — Hyalobathra Meyrick, 1885 (21 spp.) [SUTRISNO &

HORAK 2003] — Isocentris Meyrick, 1887 (7 spp.) — Pionea-

bathra J. C Shaffer & Munroe, 2007 (1 sp.) — Portentomorpha

Amsel, 1956 (1 sp.)

Pyraustini: Achyra Guenée, 1849 (19 spp.) — Adoxobotys Mun-

roe, 1978 (3 spp.) — Aglaops Warren, 1892 (4 spp.) — Anamalaia

Munroe & Mutuura, 1969 (1 sp.) — Anania Hübner, 1823 (117

spp.) — Arenochroa Munroe, 1976 (1 sp.) — Aurorobotys Mun-

roe & Mutuura, 1971 (2 spp.) — Callibotys Munroe & Mutuura,

1969 (3 spp.) — Carminibotys Munroe & Mutuura, 1971 (1 sp.)

— Ceuthobotys Munroe, 1978 (1 sp.) — Chilochroma Amsel, 1956

(4 spp.) — Chilocorsia Munroe, 1964 (1 sp.) — Chilopionea Mun-

roe, 1964 (1 sp.) — Circobotys Butler, 1879 (19 spp.) — Croci-

dophora Lederer, 1863 (24 spp.) — Crypsiptya Meyrick, 1894 (8

spp.) — Cybalobotys Maes, 2001(3 spp.) — Deltobotys Munroe,

1964 (3 spp.) — Demobotys Munroe & Mutuura, 1969 (2 spp.)

— Ecpyrrhorrhoe Hübner, 1825 (12 spp.) — Epicorsia Hübner,

1818 (9 spp.) — Epiparbattia Caradja, 1925 (2 spp.) — Eumor-

phobotys Munroe & Mutuura, 1969 (2 spp.) — Fumibotys Mun-

roe, 1976 (1 sp.) — Gynenomis Munroe & Mutuura, 1968 (2 spp.)

M et al.: Phylogenetic systematics of Spilomelinae and Pyraustinae

204

— Hahncappsia Munroe, 1976 (39 spp.) — Helvibotys Munroe,

1976 (5 spp.) — Hyalorista Warren, 1892 (5 spp.) — Limbobo-

tys Munroe & Mutuura, 1970 (5 spp.) — Loxostege Hübner, 1825

(90 spp.) — Munroeodes Amsel, 1957 (4 spp.) — Nascia J. Cur-

tis, 1835 (3 spp.) — Neadeloides Klima, 1939 (2 spp.) — Neoepi-

corsia Munroe, 1964 (7 spp.) — Neohelvibotys Munroe, 1976 (9

spp.) — Nephelobotys Munroe & Mutuura, 1970 (1 sp.) — Nomis

Motschulsky, 1861 (4 spp.) — Oenobotys Munroe, 1976 (5 spp.)

— Oronomis Munroe & Mutuura, 1968 (1 sp.) — Ostrinia Hübner,

1825 (21 spp.) — Pagyda Walker, 1859 (26 spp.) — Palepicorsia

Maes, 1995 (1 sp.) — Paracorsia Marion, 1959 (1 sp.) — Para-

nomis Munroe & Mutuura, 1968 (4 spp.) — Paratalanta Meyrick,

1890 (9 spp.) — Parbattia Moore, 1888 (6 spp.) — Perispasta Zel-

ler, 1876 (1 sp.) — Placosaris Meyrick, 1897 (20 spp.) — Powysia

Maes, 2006 (1 sp.) — Prooedema Hampson, 1891 (1 sp.) — Pro-

tepicorsia Munroe, 1964 (13 spp.) — Psammotis Hübner, 1825 (8

spp.) — Pseudepicorsia Munroe, 1964 (4 spp.) — Pseudognatho-

botys Maes, 2001 (2 spp.) — Pseudopagyda Slamka, 2013 (3 spp.)

[CHEN & ZHANG 2017] — Pseudopolygrammodes Munroe &

Mutuura, 1969 (1 sp.) — Pseudopyrausta Amsel, 1956 (6 spp.) —

Pyrasia M. O. Martin, 1986 (1 sp.) — Pyrausta Schrank, 1802 (341

spp.) — Sarabotys Munroe, 1964 (2 spp.) — Sclerocona Meyrick,

1890 (1 sp.) — Sinibotys Munroe & Mutuura, 1969 (5 spp.) —

Sitochroa Hübner, 1825 (10 spp.) — Thivolleo Maes, 2006 (4 spp.)

— Thliptoceras Warren, 1890 (31 spp.) — Toxobotys Munroe &

Mutuura, 1968 (3 spp.) — Vittabotys Munroe & Mutuura, 1970

(1 sp.) — Xanthostege Munroe, 1976 (2 spp.)

Electronic Supplement Files

at http://www.senckenberg.de/arthropod-systematics

File 1: mally&al-spilomelinaephylogeny-asp2019-electronic

supplement-1.xlsx — Table S1. List of examined genitalia slides.

— 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

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Table S1. List of examined genitalia slides

Data

May 2019

Richard Mally · James E. Hayden · Christoph Neinhuis · Bjarte H Jordal · Matthias Nuss

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Following a request from the European Commission, the EFSA Panel on Plant Health performed a quantitative risk assessment of Leucinodes orbonalis (Lepidoptera: Crambidae), the eggplant fruit and shoot borer, for the EU. The assessment focused on potential pathways for entry, climatic conditions favouring establishment, spread and impact. Options for risk reduction are discussed but effectiveness was not quantified. L. orbonalis is a key pest of eggplant (aubergine/brinjal) in the Indian subcontinent and occurs throughout most of southern Asia with records mostly from India and Bangladesh. The main pathway of entry is fruit of solanaceous plants, primarily exotic varieties of eggplant, Solanum melongena and turkey berry, S. torvum. The trade in both commodities from Asia is small but nevertheless dwarfs the trade in other Solanum fruits from Asia (S. aethiopicum, S. anguivi, S. virginianum, S. aculeatissimum, S. undatum). Other Solanum fruits were therefore not further assessed as potential pathways. The trade in eggplant from Asia consists of special fruit types and caters mostly to niche markets in the EU, while most eggplant consumed in Europe is produced in southern European and northern African countries, where L. orbonalis does not occur. Using expert knowledge elicitation (EKE) and pathway modelling, the Panel estimated that approximately 3–670 infested fruit (90% certainty range, CR) of S. melongena or fruit bunches of S. torvum enter into regions of the EU that are suitable for L. orbonalis establishment each year. Based on CLIMEX modelling, and using two possible thresholds of ecoclimatic index (EI) to indicate uncertainty in establishment potential, climates favouring establishment occur mostly in southern Europe, where, based on human population, approximately 14% of the imported produce is distributed across NUTS2 regions where EI ≥ 30; or 23% of the produce is distributed where EI ≥ 15. Escape of adult moths occurs mostly from consumer waste. By analysing results of different scenarios for the proportion of S. melongena and S. torvum in the trade, and considering uncertainties in the climatic suitability of southern Europe, adult moth emergence in areas suitable for establishment is expected to vary between 84 individuals per year and one individual per 40 years (based on 90% CR in different scenarios). In the baseline scenario, 25% of the solanaceous fruit from Asia is S. torvum, 75% is S. melongena and EI ≥ 30 is required for establishment. After accounting for the chances of mating, host finding and establishment, the probability of a mated female establishing a founder population in the EU is less than 1 in 100,000 to about 1 event per 622 years (90% CR in baseline scenario). The waiting time until the first establishment is then 622 to more than 100,000 years (CR). If such a founder population were established, the moth is estimated to spread at a rate of 0.65–7.0 km per year after a lag phase of 5–92 years. The impact of the insect on the production of eggplant is estimated to be 0.67%–13% (CR) if growers take no specific action against the insect and 0.13%–1.9% if they do take targeted actions. Tomato (S. lycopersicum) and potato (S. tuberosum) are hosts of L. orbonalis, but the insect does not develop to maturity in tomato fruit, and it does not feed on potato tubers under field conditions; hence, damage to potato can only occur due to feeding on shoots. Tomato and potato are not preferred hosts; nevertheless, impact can occur if populations of L. orbonalis are high and preferred hosts are not available. The Panel did not assess this damage due to insufficient information.

Description of two new species in the genus Glyphodes Guene from Indonesia (Lepidoptera: Crambidae, Spilomelinae)

Article

Jan 2024

Two new species of Glyphodes Guenée, 1854 from Indonesia are proposed as new to science, namely G. nurfitriae sp. nov. and G. ahsanae sp. nov. The total number of recorded Glyphodes for Indonesia is 48 at present. Images of adults and genitalia are provided for both new species.

CAN GONDWANA VICARIANCE, RATHER THAN CENOZOIC CHANCE DISPERSAL, BETTER EXPLAIN THE DISJUNCTION OF AMBULICINI HAWKMOTHS BETWEEN MADAGASCAR AND NEW CALEDONIA? Can Gondwana vicariance, rather than Cenozoic chance dispersal, better explain the disjunction of Ambulycini hawkmoths between Madagascar and New Caledonia?

Article

Full-text available

Jun 2024

The geographic disjunction between the hawkmoth genera Compsulyx in New Caledonia and Batocnema in East Africa-Madagascar is a biogeographic pattern shared with other plant and animal taxa, irrespective of their individual means of dispersal. The disjunction is almost identical to a sister species relationship in the plant genus Acridocarpus (Malpighiaceae), similar to Cunonia (Cunoniaceae) in New Caledonia and South Africa, and comparable to Dietes (Iridaceae) on Lord Howe Island and Madagascar/East Africa. These and other geographic disjunctions represent scattered elements of the numerous trans-Indian Ocean connections involving more widespread taxa, some with distributions that include the Pacific. The biogeographic patterns are consistent with Mesozoic tectonic events that disrupted the continuity of widespread ancestral distributions. The tectonic correlations are not in conflict with younger fossil or island calibrated molecular divergence ages that represent minimum estimates only. And they provide a potential falsification of centres of origin and chance dispersal models generated by ancestral-areas programmes.

Complete Mitochondrial Genome of the Eggplant Fruit and Shoot Borer, Leucinodes orbonalis Guenée (Lepidoptera: Crambidae), and Comparison with Other Pyraloid Moths

Article

Full-text available

Mar 2024

The eggplant fruit and shoot borer (EFSB) (Leucinodes orbonalis Guenée) is a devastating lepidopteran pest of eggplant (Solanum melongena L.) in the Philippines. Management of an insect pest like the EFSB requires an understanding of its biology, evolution, and adaptations. Genomic resources provide a starting point for understanding EFSB biology, as the resources can be used for phylogenetics and population structure studies. To date, genomic resources are scarce for EFSB; thus, this study generated its complete mitochondrial genome (mitogenome). The circular mitogenome is 15,244 bp-long. It contains 37 genes, namely 13 protein-coding, 22 tRNA, and 2 rRNA genes, and has conserved noncoding regions, motifs, and gene syntenies characteristic of lepidopteran mitogenomes. Some protein-coding genes start and end with non-canonical codons. The tRNA genes exhibit a conserved cloverleaf structure, with the exception in trnS1. Partitioned phylogenetic analysis using 72 pyraloids generated highly supported maximum likelihood and Bayesian inference trees revealing expected basal splits between Crambidae and Pyralidae, and Spilomelinae and Pyraustinae. Spilomelinae was recovered to be paraphyletic, with the EFSB robustly placed before the split of Spilomelinae and Pyraustinae. Overall, the EFSB mitogenome resource will be useful for delineations within Spilomelinae and population structure analysis.

The genome sequence of the Chalk Hill Pearl moth Mecyna flavalis (Denis & Schiffermüller, 1775)

Article

Mar 2024

We present a genome assembly from an individual female Mecyna flavalis (Chalk Hill Pearl moth; Arthropoda; Insecta; Lepidoptera; Crambidae). The genome sequence is 531.6 megabases in span. Most of the assembly is scaffolded into 32 chromosomal pseudomolecules, including the Z and W sex chromosomes. The mitochondrial genome has also been assembled and is 15.31 kilobases in length. Gene annotation of this assembly on Ensembl identified 12,508 protein coding genes.

Revision of the genus Marwitzia Gaede,1917 (Lepidoptera, Pyraloidea, Crambidae, Spilomelinae)

Article

Full-text available

Sep 2023

Koen Maes

Revision of the genus Marwitzia Gaede,1917 with the description of a new species.

Revival of genus Patania Moore and reporting of a new species menoni (Pyraustinar: Pyralidae: Lepidoptera)

Article

Full-text available

Feb 2020

Jagbir Singh Kirti

Phylogeny, character evolution and tribal classification in Crambinae and Scopariinae (Lepidoptera, Crambidae)

Article

Full-text available

Oct 2019

Crambinae (2047 spp.) and Scopariinae (577 spp.) are two major groups of pyraloid moths with a worldwide distribution. Their larvae feed predominantly on Poales and Bryophyta, with many cereal crop pests. We present the first molecular phylogeny of the two groups based on five nuclear genes and one mitochondrial gene (total = 4713 bp) sampled for 58 crambine species representing 56 genera and all tribes, 33 scopariine species representing 12 genera, and species in several other crambid lineages. Maximum likelihood and Bayesian analyses of the molecular data resolve suprageneric relationships in Crambinae and Scopariinae, whereas relationships between these and other subfamilies remain ambiguous. Crambinae and Scopariinae are each recovered as monophyletic groups, and Erupini, formerly regarded as an ingroup of Midilinae, is recovered as a possible sister group of Crambinae. The tree topology suggests the following two major changes within Crambinae: Prionapterygini Landry syn.n. of Ancylolomiini Ragonot stat. rev. and Myelobiini Minet syn.n. of Chiloini Heinemann. Argyriini Munroe is monophyletic after the transfer of Pseudocatharylla Bleszynski and Vaxi Bleszynski to Calamotrophini. Crambini, Diptychophorini and Haimbachiini are monophyletic after the exclusion of Ancylolomia Hübner, Euchromius Guenée, Micrelephas Dognin and Miyakea Marumo from Crambini, as well as Microchilo Okano from Diptychophorini. Euchromiini tribe n. is described for Euchromius. Microcramboides Bleszynski syn.n. and Tortriculladia Bleszynski syn.n. are synonymized with Microcrambus Bleszynski. In Scopariinae, Caradjaina Leraut syn.n. and Cholius Guenée syn.n. are synonymized with Scoparia Haworth, and, in addition, Dasyscopa Meyrick syn.n., Dipleurinodes Leraut syn.n. and Eudipleurina Leraut syn.n. are synonymized with Eudonia Billberg. Micraglossa melanoxantha (Turner) (Scoparia) comb.n. is proposed as a new combination. We analysed 27 morphological characters of wing venation, tympanal organs, male and female genitalia, as well as host plant data and egg‐laying behaviour. The ancestral character‐state reconstructions confirmed previous apomorphies and highlighted new apomorphies for some of the newly recovered clades. The derived, nonadhesive egg‐dropping behaviour is found to have evolved at least twice in Crambinae and is associated with the use of Pooideae as host plants. This published work has been registered in ZooBank, http://zoobank.org/urn:lsid:zoobank.org:pub:1A84282D‐930A‐4C32‐8340‐D681BFF27A12. The Crambinae + Scopariinae phylogeny is reconstructed with five nuclear genes and one mitochondrial gene (total = 4713 bp) sampled for 91 taxa representing 68 genera. The tribal classification in Crambinae is revised and Euchromiini tribe n. is described, and in Scopariinae, five genera found nested within the species‐rich Eudonia and Scoparia are synonymized. The behaviour of dropping nonadhesive eggs is found to have evolved at least twice in Crambinae and is associated with the use of Pooideae as host plants.

Molecular Phylogeny of the Indo-Australian Glyphodes and its Allied Genera (Insecta: Lepidoptera: Crambidae: Spilomelinae) Inferred from Mitochondrial COI and COII and Nuclear EF-1α Gene Sequences

Article

Full-text available

Feb 2006

The phylogeny of Indo-Australian Glyphodes (15 species), Talanga (three species), Agrioglypta (four species), and two outgroup species, Ostrinia nubilalis Hubner, 1825 and Metallarcha aureodiscalis (Hampson, 1918), was inferred from nucleotide sequence variation in the mitochondrial COI (686bp) and COII (687bp) genes and the nuclear EF-1α (973bp) gene. While the third codon positions in COI and EF-1α were not saturated with substitutions, those in COII were saturated when sequences were compared between genera. MP analyses based on pooled data of COI, COII, and EF-1α resulted in two MP trees. The strict consensus tree showed that Glyphodes, Talanga, and Agrioglypta were all not monophyletic. Another analysis based on pooled data of all molecular and morphological data resulted in a single tree, in which monophyly of the genus Glyphodes was barely supported (bootstrap value <50%). Glyphodes species were divided into the same three clades as in an earlier published, morphology-based tree, one of these clades (the so-called Group 3) being supported by a bootstrap value of 100%. In all analyses the genus Agrioglypta was shown to be paraphyletic, since A. eurytusalis (Walker, 1859) fell within the genus Talanga.

A new Indian species of shoot and capsule borer of the genus Conogethes (Lepidoptera: Crambidae), feeding on cardamom

Article

Full-text available

Jan 2018

A new species, Conogethes sahyadriensis sp. nov. (Lepidoptera: Crambidae), feeding on cardamom, is described from India. The species status is supported by diagnostic morphology as well as by genetic data. A phylogenetic analysis based on the publicly available Conogethes COI barcode sequences finds C. sahyadriensis as sister to C. pluto, and it further reveals a number of clades that potentially represent additional undescribed species.The new species is delineated from closely related and superficially similar species of Conogethes.

On the taxonomy of the erythrina moths Agathodes and Terastia (Crambidae: Spilomelinae): Two different patterns of haplotype divergence and a new species of Terastia.

Article

Full-text available

Jan 2015

Members of two species complexes in the family Crambidae feed on ornamental and medicinal plants of the genus Erythrina (Fabaceae), commonly known as coral beans and/or coral trees, in tropical and subtropical regions worldwide. Erythrina leaf-rollers (genus Agathodes) and erythrina borers (genus Terastia), although different ecologically and phenotypically, are sister genera. Each is represented by several very similar species. The erythrina leaf-rollers comprise a species complex that, in the New World, consists of Agathodes monstralis Guenée, 1854 rev. stat. and A. designalis Guenée, 1854. We analyzed morphology and DNA barcoding data of populations throughout the geographic range and concluded that North American A. monstralis, which was previously demoted to the status of a subspecies of A. designalis, is a valid species. The two are sympatric in Costa Rica. In contrast, the erythrina borer, Terastia meticulosalis Guenée, 1854, which has a similar geographic distribution to A. designalis, shows no divergence in DNA barcodes and morphology in the New World. However, DNA barcode analyses and detailed morphological examinations of the Asian and African populations of Terastia meticulosalis species group show that these represent several species. Thus, a new species, Terastia africana Sourakov, sp. nov., is described and the status of T. subjectalis Lederer, 1863 as a valid species is confirmed.

Revision of the Australian species of Agrotera Schrank (Lepidoptera: Pyraloidea: Crambidae: Spilomelinae)

Article

Full-text available

Dec 2017

The genus Agrotera Schrank, 1802 is revised for Australia and the generic definition is refined based on the male genitalia. The genera Leucinodella Strand, 1918 stat. rev. with L. leucostola (Hampson, 1896) comb. nov., Nistra Walker, 1859 stat. rev. with N. coelatalis Walker, 1859 comb. rev., Sagariphora Meyrick, 1894 stat. rev. with S. magnificalis (Hampson, 1893) comb. nov., and Tetracona Meyrick, 1884 stat. rev. with T. amathealis (Walker, 1859) comb. rev. and T. pictalis Warren, 1896 comb. rev. are removed from synonymy with Agrotera, as they lack the synapomorphies of Agrotera. Two new species, Agrotera genuflexa sp. nov. from Northern Territory, Queensland and New South Wales, and A. longitabulata sp. nov. from Queensland, are described. The taxonomic status of the Australian species of Agrotera is discussed, and a key to all species, based on males, is provided. The adults and genitalia of the new species and some related species are figured.

Variably hungry caterpillars: Predictive models and foliar chemistry suggest how to eat a rainforest

Article

Full-text available

Nov 2017
Proc Biol Sci

A long-term goal in evolutionary ecology is to explain the incredible diversity of insect herbivores and patterns of host plant use in speciose groups like tropical Lepidoptera. Here, we used standardized food-web data, multigene phylogenies of both trophic levels and plant chemistry data to model interactions between Lepidoptera larvae (caterpillars) from two lineages (Geometridae and Pyraloidea) and plants in a species-rich lowland rainforest in New Guinea. Model parameters were used to make and test blind predictions for two hectares of an exhaustively sampled forest. For pyraloids, we relied on phylogeny alone and predicted 54% of species-level interactions, translating to 79% of all trophic links for individual insects, by sampling insects from only 15% of local woody plant diversity. The phylogenetic distribution of host-plant associations in polyphagous geometrids was less conserved, reducing accuracy. In a truly quantitative food web, only 40% of pair-wise interactions were described correctly in geometrids. Polyphenol oxidative activity (but not protein precipitation capacity) was important for understanding the occurrence of geometrids (but not pyraloids) across their hosts. When both foliar chemistry and plant phylogeny were included, we predicted geometrid-plant occurrence with 89% concordance. Such models help to test macroevolutionary hypotheses at the community level.

Lathrotelidae Clarke, 1971: a rehabilitated name deserving subfamily rank (Lepidoptera, Crambidae)

Article

Jan 2015

Joël Minet

New pyralidae (lepidoptera) from the papuan region. III

Article

Jan 1974

E. Munroe

The following new Pyraustinae are described: Goliathodes n. gen., type-species G. shafferi n. sp., New Guinea and Waigeu; Spoladea mimetica n. sp., New Guinea; Meekiaria n. gen., type-species M. lignea n. sp., New Guinea; Aiyura n. gen., type-species A. linoptera n. sp., New Guinea.

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

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