MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 43 A WORLD REVISION OF THE GENUS SPODOPTERA GUENEE (LEPIDOPTERA: NOCTUIDAE) By MICHAEL G. POGUE ty eet Vy, a “np v “ing Ming YY > TNS Mh \\\ TTA PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 2002 The American Entomological Society Philadelphia Founded 1859 Incorporated 1862 PUBLICATIONS COMMITTEE — Howard Boyd, Chairman Daniel Otte Norman E. Woodley The offices, library, and meeting rooms of the Society are at The Academy of Natural Sciences of Philadelphia PUBLICATIONS OF THE SOCIETY TRANSACTIONS OF THE AMERICAN ENTOMOLOGICAL Society are published in annual volumes of quarterly numbers. Subscription prices: United States $20.00 per yearly volume (four issues); foreign countries $22.00 per volume; U.S. and foreign members of the society $10.00 per volume. Entomotocicat News is published bimonthly except July-August. Subscription prices: $20.00 per annual volume; U.S. and foreign members of the society $10.00 per volume. Memoirs oF THE American Entomotocica Society are published irregularly when a suit- able monograph is submitted and accepted for publication. Prices vary with size; see back cover for list of available numbers. Please address all inquiries relative to publications and other society matters to: The American Entomological Society The Academy of Natural Sciences 1900 Benjamin Franklin Parkway Philadelphia, PA 19103 USA e-mail: AES@say.acnatsci.org Please visit the society’s internet website at: http:/ /www.acnatsci.org /aes/ MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 43 A WORLD REVISION OF THE GENUS SPODOPTERA GUENEE (LEPIDOPTERA: NOCTUIDAE) By MICHAEL G. POGUE WNUUUUL ECCT yy Minry ‘My TOMO; SK PUBLISHED BY THE AMERICAN ENTOMOLOGICAL SOCIETY AT THE ACADEMY OF NATURAL SCIENCES PHILADELPHIA 2002 Norman E. Woodley Editor Issued 28 January 2002 PRINTED IN THE UNITED STATES OF AMERICA by Sheridan Books Chelsea, Michigan TABLE OF CONTENTS IIMIRICKUETOM oosobcocecassvsosnesocc0b0g 00s deHd oOo OadEnobODOSUaGonDDOOOOD AL 1 lEtrarelghne NOVO comanaepondsoe ooo eb eS eS tA piomen cacao eels amen Amt SAL 2 Wanvalilostielamts meyers ct sens ate servers seer sial sete svevnis exerorers a loieislareconeuatelete: ain sieneds 3 INfethod sand fatertall Simererrr rere ora tente pre oiekcearterod re areease eeraceeees 4 DESC OIONS. coccpoo de cons son sono cau anus ounce Dodo oUmeoeeuos Ba Maesds c 4 SourcesiotsMateriallE'xaminedsrmrse ccc cman cri tix: ce cies seem ae ce eee 4 IMEVOOINE: o soa babdoubaoc noone at obEDgsEaoboCodeS CUD OE DoS EMU Se soins Memmce 5 Databasesaresrpaverpecrcusvecraveitesaucneuexcieohssneyecesgevereusitoue suenersPaccl gene veces Sessa aueyhsfanapweatiacens 5 Manvalltlostiblantsteevwrere ci niun seit ttcarrctcittalertiele cree ciclsie are inlevan etiee 5 Naturally eme miles weir aryerncye cus sige mcancacicl-e rasta tii iovp urns nes, ceteris ae ecisetgels nk 5 Specimeniareparationeacem ccc ny eke een nies / tse i-iae isesreee ree 6 Morphologicalalermspen sn cmnerre ore riser cere a iene nym iene ed rese va renyerae 7 GladisticyAmalySisy ener sk tvciccect Backers terre auctor austen c ceca spe tahite) Resi sosist teers 8 @lacIStHGIReSUltsyvavarart tec usreyytecsrey space ciel axertoreuevene aes atecsee ats selitescallascedelie avevslaonieeene 9 Phylogenetic Gharacterswenna eect Sai cerca ois seracr isla) asi 12 IBISEIOly Seman Bionic pala casi nea Dee Oe: Ceo Re RECO Ree ro TS ic ct se och a Pre 14 PUM reaxgehar tee rte cis Sheetal esas ster ate aeakc wean terra iar tr saeai ois sfenansie vive Seale ganeite wile 15 MialetGemitaliaaearrersct ck attra een cuyrr ttt Morey spps petals staves ase vorelaye fale sae 17 emale:Gemitaliatateyesrcter aca yrsrees i aatesoncpsicecimneiniey tae ere) sks 5 oie sysimoik ste) speyeneveliele = 31 Acuiltsystemati chlireatmentre cere neeUern ae oe Creer Tie sree eae nny 35 Ge Ch IS tee Grater tesctreere rete tapta ster sels pares Wieceneraestons i sae el aie secaranaitedern auaatoter sé 35 Spodoptera GUeMee rpc yyectae oye rN nMn Leo eer tal nastier nse lay alelbsiona evn el 36 INCViSLOI SPECIES a rey raaeetret ey Usp ce ee eal lave a a natal Spits op Maite hoe eter g Nievede sue 37 Key toyNdultsiofi@ldiWorldiSpeciesinn ssaivsiiscal relay feisusss secs sp aycteseb sry eet = 37 Key to Adults of New World Species ................ 000 c eee e eee eee eee 38 ANohulhs |DisxedhoO mnaocaccangodasseokenedt oor eamorace Dh aU EE suo mmacomemas 40 Eanvalisystematicilreatmentirn. teenie ition ics ane ee) seers 97 XO SAU HAGILANED soocdocodsscnonssounsdvenonoosanobonsenddabEEeEae 98 Warviale Deserip ti Orsi ye crveys sence hes hells ouselouedsncusveususketeqsys icy Sieiaysrseswseisaecsuelovepsns)itare 99 ENG ANONAIS (exaaTA ONS. Noe aH oan Soa aoo pe oto ote une om OCuros hou Oona Ratan ome Here 116 Peiteratwres Gite diy yreuscyorre vtec sete ce tesece Roe votoneeeycisi sacs eusfade ssere sion tseyew custo bertvanobeterels onsyehe AZ, AND OLUCIE RI yao 6 ope vaca we oe ae Oote he OARS Coon Orr Br ore Dim raniiirl Emit mura eer 125 PAP PEM IX AM Hover cece cehecueee ckctarekar eyes ou steney heh ste ees ou ant cul yusua falas tspay saunas ae bat anday sbabaliscavaubs 126 /NDVOLIMCID SIND, soe aterbiwerS alee See alah SHORTS coatns Clair Alerunchie renin. aio iibal Saar men much ee 127 | atbeqbanet) IVA byHehriin ao Grates alo cram wrya aid roe oat Ot cn IIIS Sia Section Gcaee MON Aer R Oar 128 IGgYSKED-€ ola Gietore: Sa cert aleve sas tice tire ech ean areas Oe nn nr SO ERR Pe 201 a fhe , 7 mary i Fi LMS ni ai ‘ i 1 MEMOIRS OF THE AMERICAN ENTOMOLOGICAL SOCIETY NUMBER 43 A World Revision of the Genus Spodoptera Guenée (Lepidoptera: Noctuidae) MICHAEL G. POGUE Systematic Entomology Laboratory, ARS, USDA Smithsonian Institution NHB-168 Washington, DC 20560-0168 USA e-mail: mpogue@sel.barc.usda.gov ABSTRACT—The world fauna of the armyworm genus Spodoptera Guenée is revised. A phylogeny of Spodoptera was construct- ed based on cladistic relationships of the 30 species. Two genera that are sister groups of Spodoptera based on a molecular phy- logeny of the Noctuidae served as outgroups. The data set contained 24 morphological characters (6 binary and 18 multistate) comprised of 134 character states. Cladistic analysis used the parsimony program Hennig86 which constructed two trees with a length of 143, a consistency index of 0.77, and a retention index of 0.91. Eight monophyletic clades containing 2 to 5 species were recognized. Two species were not part of any of the eight clades, but were sister taxa to larger monophyletic groups on the tree. These were S. exigua (Hiibner) which was the most plesiomorphic species and S. frugiperda (J.E. Smith) was the sister species to the dolichos, littoralis, evanida, and praefica clades. Descriptions, distributions, and illustrations of all adults are included. Twenty species of larvae are described and 17 are illustrated. Keys are provided to adults based on geographical distribution, external morphology, and male and female genitalia. A key to larvae is based on color pattern and morphology. Ten neotypes and 32 lec- totypes are designated and 5 new synonyms are proposed. Specimens examined during this study were given a bar code label. Databases of the bar coded specimens and all unique localities encountered during this study were developed. Databases were produced for larval host plants, parasitoids, predators, and a bibliography to the records contained in these databases. Databas- es can be found on the internet at the Systematic Entomology Laboratory and the Department of Entomology, Smithsonian In- stitution websites. INTRODUCTION Some of the most serious economic armyworm pests are species in the genus Spodoptera Guenée. Species are polyphagous on numerous vegetables, grain crops, and pasture lands. The genus is main- ly tropical and subtropical throughout the world with some species migrating into more temperate regions throughout the growing season, e.g., Spo- doptera exigua (Hubner) and S. littoralis (Boisduval) have been collected in northwestern Europe (Mikkola & Salmensuu 1965, Goater 1983). In the Western Hemisphere those species that are found in the temperate regions of the United States in- clude S. exigua, S. frugiperda (J. E. Smith), S. or- nithogalli (Guenée), S. praefica (Grote), S. eridania (Walker), and S. latifascia (Walker). In the temper- ate regions of South America S. frugiperda and S. cosmiodes (Walker) are pests. About half of the 30 Spodoptera species have been considered pests. No phylogeny of the genus has ever been constructed using modern cladistic techniques. To better understand the re- lationships between pest and non-pest species of Spodoptera and possible host plant relationships, a phylogenetic analysis of the genus is presented. N No comprehensive revision of Spodoptera on a world level is available. Brown & Dewhurst (1975) treated the African species of Spodoptera, giving keys to adults, larvae, and some pupae. They reviewed the taxonomic literature concern- ing generic synonyms of Spodoptera and the sepa- ration of S. littoralis from S. litura (Fabricius), dis- cussed character states used to separate both species of adults and larvae, and gave a list of common names used in Africa for the economi- cally important species. Larval host plants were discussed as well as species distributions. Todd & Poole (1980) treated the Western Hemisphere species of Spodoptera by providing a key to the adults, and illustrating the adult and male and fe- male genitalia. These works cover 22 of the 30 species, but the other 8 species, including eco- nomically important species such as S. litura and S. depravata (Butler), have not been included in any faunal or revisionary work on Spodoptera. Spodoptera descoinsi Lalanne-Cassou & Silvain (Lalanne-Cassou & Silvain in Lalanne-Cassou, et al. 1994) was described from French Guiana and is closely related to S. latifascia and S. cosmiodes. Spo- doptera cosmiodes was recently found to be a valid species and not a synonym of S. latifascia (Silvain & Lalanne-Cassou 1997). All three species are dis- tinct in genitalic structures and pheromone com- position. The phylogeny presented in this paper was constructed using the parsimony program Hen- nig86, Version 1.5 (Farris 1988). A total of 24 char- acters were used including 6 binary and 18 multi- state. Transformation series analysis (TSA) was used to construct character state trees of the mul- tistate characters (Mickevich 1982). Character state trees were used to analyze 14 of the multi- state characters. Four of the multistate characters were run ordered. Eight monophyletic clades con- taining 2 to 5 species were recognized. Spodoptera exigua was the most plesiomorphic and S. fru- giperda was the sister species to the dolichos, lit- toralis, evanida, and praefica clades. Two outgroups were used based on results of a molecular system- atics study of the Noctuidae (Mitchell 1998). A single tree was produced with a length of 143, a consistency index of 0.77, and a retention index of 0.91 (Fig. 3). Databases were produced for the larval host plants, parasitoids, and predators of Spodoptera, as well as a bibliographic database in support of these. A specimen database was created for exam- REVISION OF SPODOPTERA GUENEE ined material. A gazetteer is provided for all local- ities recorded in this revision, including latitude and longtiude data when available. These are available on the internet at the Systematic Ento- mology Laboratory and the Department of Ento- mology, Smithsonian Institution websites. Todd & Poole (1980) transferred Spodoptera hip- paris (Druce) out of Leucochlaena. This genus is re- stricted to the Old World and its species superfi- cially resemble S. hipparis, but the male and female genitalia of S. hipparis indicate that it is not congeneric with species of Leucochlaena. Spodop- tera hipparis should not be included in Spodoptera because it shares no synapomorphies with Spo- doptera, such as the large hair tuft on the female 8th segment. A review of New and Old World genera should be conducted to determine where S. hipparis should be placed. If no satisfactory generic placement can be found, a new one will be described to accommodate this species. LITERATURE REVIEW Spodoptera was described by Guenée (1852a) and included the following 7 species: S. mauritia (Boisduval), S. abyssinia (Guenée) (in Ariathisa (Poole 1989)), S. acronyctoides (a synonym of S. mauritia), S. nubes Guenée (a synonym of S. mauri- tia), S. pecten Guenée, S. filum Guenée (a synonym of S. mauritia), and S. cilium Guenée. He also cre- ated the unnecessary objective replacement name, S. acronyctiformis Guenée (1852b), for S. acronyc- toides Guenée (Poole 1989). Three other species were originally described in Spodoptera, S. evanida Schaus (1914), S. malagasy Viette (1967) and S. de- scoinsi Lalanne-Cassou & Silvain (1994). Two synonyms of Spodoptera, Laphygma and Prodenia were described by Guenée (1852a). La- phygma was synonymized by Turner (1920) and Prodenia was synonymized by Viette (1963). Gue- née described the species cycloides (a synonym of cillum) and macra (a synonym of frugiperda), and placed the species exigua, pygmae (a synonym of exigua), and frugiperda in Laphygma. He described the species tasmanica (a synonym of litura), or- nithogalli, ciligera (a synonym of litura), eudiopta (a synonym of ornithogalli), and testaceoides (a syn- onym of littoralis), and placed the species androgea, commelinae (a synonym of dolichos), and retina (a synonym of littoralis) in Prodenia. Guenée (1852a) also described the genus Xy- lomyges in which he described the species sunia. M. G. POGUE 3 Hampson (1909) described ochrea and placed eri- dania, sunia, and peruviana in Xylomyges. Poole (1989) points out that Spodoptera albula (Walker) is the species that had been previously known as “Spodoptera sunia (Guenée)” however, the real Xy- lomyges sunia Guenée is actually a synonym of Neogalea esula Druce. All references relating to Spodoptera sunia prior to 1989 should be referred to Spodoptera albula. I am not going to detail the works of Walker (1856, 1857a, 1857b, 1858, 1862a, 1862b, 1865a, 1865b, 1869, 1870) who described species of Spo- doptera, but will summarize his results. Walker created 48 names of which 8 are valid (17%). He placed these species in 10 genera and used Spo- doptera only once, in describing S. insula (a syn- onym of cilium). Druce (in Godman & Salvin 1889) listed 4 species under both Laphygma and Prodenia for the Mexican and Central American fauna. No new species were described. Hampson (1894) placed littoralis (litura) and Catamecia minima (Swinhoe) in Prodenia, and in- cluded Spodoptera mauritia and Calogramma festiva (Donovan). Spodoptera picta (Guérin-Meéneville) is the subjective replacement name of Calogramma festiva, a junior primary homonym of Phalaena fes- tiva Cramer (1775) (Sugi 1982). He also described two synonyms of Spodoptera apertura (Walker), Prodenia synstictis (Hampson 1896) and Prodenia leucophlebia (Hampson 1902). Hampson (1909) dealt with all of the Spodoptera species available to him at the time. He described rubrifusa (a syn- onym of androgea), and placed dolichos, pulchella, androgea, litura, latifascia, ornithogalli, praefica, and marima in Prodenia. He placed umbraculata, pecten, Ariathisa abyssinia, compta, mauritia, and Diarsia postfusca in Spodoptera. He placed leucophlebia (a synonym of apertura), apertura, exempta, frugiperda, flavimaculata (a synonym of exigua), and exigua in Laphygma. In the genus Xylomyges, he included eridania, peruviana (synonym of ochrea), ochrea, and sunia (albula). He also included C. festiva. Turner (1920) gave descriptions of the Spodop- tera species in Australia. He listed Calogramma fes- tiva (S. picta), Prodenia litura, and the following Spodoptera species: umbraculata, mauritia, leuco- phlebia (=apertura), exempta, and exigua. Edwards (1996) lists the following species of Spodoptera from Australia: picta, litura, mauritia, apertura, ex- empta, exigua, and umbraculata. Viette (1967) described and illustrated 6 species of Spodoptera from Madagascar. He de- scribed a new subspecies, S. leucophlebia malagasy. This is now recognized as a valid species, S. mala- gasy Viette (Poole 1989). Brown & Dewhurst (1975) revised the Spodop- tera species of Africa and the Near East. This is the most comprehensive treatment of Spodoptera for any region. They included descriptions of 8 species for both adults and larvae, treated 5 species of pupae and gave keys to these life stages. They discussed common names, distribu- tion, characters to aid in identification, and host plants. They included illustrations of adults, lar- vae, pupae, spermatophores, male and female genitalia as well as other characters used in the keys. Todd & Poole (1980) treated the Spodoptera species of the Western Hemisphere. They present- ed keys based on maculation and male genitalia. Fourteen species were treated, including discus- sions of characters used to identify them and their distributions. Adults and both male and female genitalia were illustrated. Holloway (1989) discussed 8 species of Spodop- tera from Borneo. He gave diagnoses, discussed their distribution, habitat preference, and biology including some host plant records. All adults were illustrated, including male genitalia, but not all female genitalia were illustrated. He listed 3 species from Mt. Kinabalu, Sabah (Holloway 1976); and the widespread pest species S. mauritia and S. litura from New Caledonia (Holloway 1979) and Norfolk Island (Holloway 1977). These pests were also listed by Robinson (1975) from Fiji. Lalanne-Cassou et al. (1994) described a new species of Spodoptera from French Guiana that is closely allied to S. latifascia. They include a re- vised couplet for the key in Todd & Poole (1980) and information on the differences between the female sex pheromone of these two species. They also resurrected from synonymy S. cosmiodes based on differences in the genital morphology, mitochondrial DNA, pheromones, and behavior (Silvain & Lalanne-Cassou 1997). LARVAL Host PLANTS The common name for Spodoptera is “army- worm”, because of the behavior of many species that move en masse from one location to another when searching for preferred hosts. Highly mo- 4 REVISION OF SPODOPTERA GUENEE bile, large larvae may have a tendency to be more polyphagous than smaller sedentary larvae (Reavey 1993). Being highly mobile gives larvae the potential of a broader range of host plants (Berdegue & Trumble 1996). All clades have at least one pest that is a serious threat to agriculture in the world, except the umbraculata clade. Certain clades (Fig. 3) can be classified with respect to host plant specificity and polyphagy. The cilium clade feeds on 10 or fewer plant species, most, but not all of which are grasses (Poaceae) and could be considered oligophagous. In the umbraculata clade larval host plants are known from only 2 species. The larva of Spodop- tera pectinicornis (Hampson) is monophagous on Pistia stratiotes (L.) (water lettuce, Araceae), and Spodoptera apertura (Walker) has been collected in the field on tobacco in Zimbabwe and larvae have been reared from it, but this seems to be an isolat- ed occurrence (Brown & Dewhurst 1975). Clade 21 is of Neotropical distribution and can be con- sidered polyphagous, with the exception of S. ochrea (Hampson). The exempta clade could be considered polyphagous and they feed almost ex- clusively on Poaceae. In this clade, S. triturata (Walker) has been recorded only on sugar cane and rice. The littoralis clade is highly polyph- agous, with the exception of S. picta. The evanida clade is oligophagous with S. cosmiodes feeding on 6 plant species and S. latifascia feeding on 8 species. The praefica clade could be considered po- lyphagous with the exception of S. pulchella (Her- rich-Schaffer) and S. marima (Schaus) (1 host spe- cies each). No biological information is known for S. roseae. Studies on host plant preference in pest species of Spodoptera have been conducted to ex- amine the preference between crop hosts. The re- sults are influenced by plants that the experi- menters choose in these studies. In India, S. exigua preferred pea (Pisum sativum L.) in one study (Rawat et al. 1969) and in another it preferred al- falfa (Medicago sativa L.) (Verma et al. 1976), but these studies are not comparable because neither of these two hosts were compared to each other. In Egypt, four studies were conducted on host preference in S. littoralis, and castor (Ricinus com- munis L.) was preferred in all of them (Elbadry et al. 1971, Abdel-Salam et al. 1971, Harakly & Bishara 1974, Aboul Nasr et al. 1975). More recent studies of host preference have focused between natural hosts and crop hosts. Pest species such as S. exigua, S. frugiperda, and S. latifascia can develop on many hosts, but optimal development may take place on non-crop plants (Berdegue & Trum- ble 1996, Portillo et al. 1996). This is important when designing intercropping scenarios for con- trol of Spodoptera pests. Weedy fields had lower numbers of S. frugiperda attacking sorghum than in weed free fields or where sorghum was inter- cropped with maize (Castro et al. 1989). METHODS AND MATERIALS DESCRIPTIONS DELTA (Descriptive Language for Taxono- my) is a computer program that enables systema- tists to gather descriptive systematic data and present it in a consistent, comparable manner (Dallwitz 1980; Dallwitz, et al. 1993). The descrip- tions for both adults and larvae were generated using DELTA. The key to the larvae was pro- duced by DELTA. Keys to the adults were not produced by DELTA, but it was used as an aid to construct these keys. SOURCES OF MATERIAL EXAMINED The following is a list of acronyms of institu- tional or private collections used during this study. Acronyms from institutional collections follow Heppner & Lamas (1982). AMNH American Museum of Natural Histo- ry, New York, New York, USA ANIC Australian National Insect Collec- tion, CSIRO, Canberra, Australia BM Private collection of Bryant Mather, Clinton, Mississippi, USA BMNH The Natural History Museum (for- merly the British Museum (Natural History)), London, England BPBM Bernice P. Bishop Museum, Honolu- lu, Hawaii, USA CNC The Canadian National Collection, Research Branch, Agriculture Cana- da, Ottawa, Canada CSU Colorado State University, Ft. Collins, Colorado, USA INBIO Instituto Nacional de Biodiversidad, Santo Domingo, Costa Rica MNHP Muséum National d’Histoire Na- turelle, Paris, France MSU Mississippi State University, Missis- sippi State, Mississippi, USA M. G. POGUE 5 ULK University of Louisville, Louisville, Kentucky, USA Department of Entomology, National Museum of Natural History, Smith- sonian Institution, Washington, D.C., USA Private collection of Vitor O. Becker, Planaltina, D.F., Brazil During the course of this study, types were searched for in the collections of the BMNH, CNC, MNHP, and USNM. If the type repository is listed as “unknown” in the type material section of the species descriptions, then it can be assumed that it is not in any of the collections that were searched. USNM VOB MAPPING The maps of collecting localities (Figs. 522— 543) were produced using the software Versamap for Windows (http://www.versamap.com). Ver- samap requires latitude and longitude in deci- mal format to plot the points on the maps. Lati- tude and longitude were obtained for collecting localities (label data) for the United States and its territories from the United States Geological Sur- vey’s (USGS) Geographic Names Information Sys- tem (http://mapping.usgs.gov/www/gnis); for the rest of the world the National Imagery and Mapping Agency’s (NIMA) GEOnet Names Serv- er was used (http://164.214.2.59/egns/html/ index.html). From these sources a Gazetteer of Spodoptera Collecting Localities database was produced which is available on the internet at the Systematic Entomology Laboratory and Depart- ment of Entomology, Smithsonian Institution websites. Localities marked “unknown” in the gazetteer were either not in the USGS and NIMA databases, or there were multiple localities with the same name, so it was impossible to infer the correct locality from the inadequate label data. DATABASES In support of this research on the world species of Spodoptera several databases were developed. Databases include those of larval plant hosts, par- asitoids, and predators including a bibliographic database that supports the records found in these databases. Not all specimens examined were indi- vidually databased, but each locality from which specimens were examined is included in the Spec- imen Database. Specimens from the BMNH and MNHP were not individually databased as their holdings were too great. Specimens examined from the other institutions were databased and each specimen was given a unique number (bar code label or individual number label), and all la- bel data, species identification, and repository where specimen is housed were recorded. A Gaz- etteer of Spodoptera Collecting Localities database was produced that includes the latitude and longi- tude of all collecting localities of Spodoptera that were examined during this study. All of these databases can be found on the internet at the Sys- tematic Entomology Laboratory and Department of Entomology, Smithsonian Institution websites. Larval Host Plants The host plant data was compiled from the lit- erature and specimens in collections. The Review of Applied Entomology, Series A, Agricultural, vol- umes 1 to 82 (1913-1994) was used to obtain most of the host plant records. A host plant record con- sists of the host plant, the Spodoptera species feed- ing on the host, and the country. The state was used for records in the United States. I tried to use the first time the record was mentioned in the lit- erature. Many older records are based on obscure Agriculture reports and those of country and/or state Entomologists. I tried to duplicate these records by more recent, easily retrievable liter- ature. Since Spodoptera are easily reared in the laboratory on a variety of hosts and artificial di- ets, many of the more common pests have been used as experimental animals. These records were ignored as the larvae may never have a chance to encounter what they were being fed in the lab- oratory in nature. I tried to base the larval host plant list on natural occurring records. All names were checked against the U. S. Department of Agriculture, Agricultural Research Service GRIN taxonomic database (www.ars-grin.gov/npgs/ taxgenform.html) which provides the structure and nomenclature for the accessions of the Na- tional Germplasm System. Names were also checked in Index Kewensis. Natural Enemies Natural enemies refer to both parasitoids and predators of Spodoptera. The primary function of natural enemies from an agricultural perspective is that of biological control of pest species. Thompson (1946 a, b, c) compiled a list of the par- asitoids of insect pests. Thompson & Simmonds 6 REVISION OF SPODOPTERA GUENEE (1964) compiled a list of insect predators. The records for these lists were compiled from the pa- pers in the Review of Applied Entomology, Series A, Agricultural, volumes 1-25 (1913-1937). I contin- ued compiling Spodoptera records from volumes 26-82 (1938-1994). A parasitoid or predator record consists of the natural enemy, the Spodop- tera species preyed upon, and the country. The state was used for records in the United States. I tried to use the first time the record was men- tioned in the literature to keep redundancy to a minimum. Some records will have more than one citation if the older record is from an obscure pub- lication, such as yearly reports from country and/or state agricultural departments and ento- mologists. Since several species of Spodoptera are easily reared, they have been used in laboratory experiments. I did not include records of para- sitoids or predators being fed Spodoptera larvae when conducting compatibility studies, mass rearings in conjunction with biological control re- leases, or experiments on host preference. The information presented should be used with caution and original papers should be con- sulted because their factual value should be eval- uated. As mentioned in Thompson & Simmonds (1964), some of these records are erroneous, how- ever excluding such references is not justified, since this is a compilation of published records. Parasitoids The nomenclature of the parasitoids was checked against the most recent catalogs and cor- rected from the original papers. Catalogs used were Krombein et al. (1979) for Hymenoptera and Stone et al. (1965), Guimaraes (1971), Delfinado & Hardy (1977), Crosskey (1980), and Evenhuis (1989) for Diptera. Nomenclature of Spodoptera was also corrected from the original papers. For example, S. litura from Africa should be referred to as S. littoralis. Many parasitoids have been re- leased from their country of origin for use in bio- logical control in other countries. A record in- cludes the Spodoptera host species, the parasitoid, and the country and/or state in the United States where the parasitoid was collected. The records are for specimens that were reared from field col- lected eggs, larvae, or pupae of Spodoptera species. Predators The nomenclature of the predators and Spo- doptera species was updated from modern cata- logues and corrected from the original papers. Catalogues used were Lorenz (1998 a, b) for the Carabidae (Coleoptera) and Henry & Froeschner (1988) and Aukema & Rieger (1996 a, b) for the Heteroptera. A record includes the Spodoptera host species, the predator, and the country and/or state that the predator was collected. The records are for specimens that were observed feeding on either eggs, larvae, or pupae of Spodoptera and were field collected. SPECIMEN PREPARATION Whole body dissections and wing prepara- tions were done by removing the left pair of wings and mounting them with white glue to a rectangular piece of acid free label paper and pin- ning them with the labels from the specimen. The right pair of wings were soaked in a 10% solution of hydrochloric acid for a few minutes then trans- ferred to bleach until the wing scales turned white. The scales were then carefully removed from both surfaces of the wings and then trans- ferred to acid fuchsin stain in 95% ethyl alcohol for 24 hours. The wings were then removed from the stain and rinsed in 95% ethyl alcohol. They were then transferred to xylene for 1 hour and mounted in Canada Balsam on a microscope slide with an appropriately sized cover slip. The whole body was put in 10% potassium hydroxide (KOH) in a test tube and heated for 30-40 minutes using a dry bath. The temperature was set so the specimen did not boil. The specimen was then cleaned and rinsed in distilled water and placed in a vial of 40% ethyl alcohol for study. After study, the whole body was placed in a 1/2 dram shell vial containing glycerine and permanently stored with the left pair of wings. To prepare the genitalia the abdomen was bro- ken off and placed in 95% ethyl alcohol to wet it, then it was placed in a test tube of 10% KOH and heated for 30-40 minutes (depending on size) us- ing a dry bath. By first wetting the abdomen in al- cohol it does not have a tendency to float in the KOH. The abdomen was then transferred to dis- tilled water and cleaned of internal matter and scales. The abdominal pelt with genitalia attached was then placed in a solution of chlorazol black and distilled water until stained, (usually a few minutes depending on strength of stain), rinsed in distilled water, and then the genitalia were dis- sected from the abdomen. The abdominal pelt was slit up the right side with ventral side up and M. G. POGUE 7 any additional cleaning was done. The genitalia and abdomen were then stained in mer- curochrome and rinsed in distilled water. The ae- doeagus was dissected from the male genitalia and the vesica was everted with distilled water using a number 30 needle on a hypodermic sy- ringe. The aedoeagus was then placed back in the chlorazol black until the vesica was lightly stained. After staining, the vesica was rinsed by shooting a stream of 99% isopropyl alcohol through it and then it was stored in isopropyl al- cohol until slide mounting. The isopropyl alcohol hardens the vesica to help hold its shape. The ab- domen and genitalia were then soaked in increas- ing strengths of 40%, 70%, and 95% ethyl alcohol for at least 2 hours each. They were then soaked in xylene for 1 hour and mounted in Canada Bal- sam. The abdomen was mounted under a sepa- rate cover slip from that of the genitalia. In some preparations the right valve of the male genitalia was removed and mounted under a separate cov- er slip. The same technique applies to the female. The coverslip over the male and female genitalia was supported by small thicknesses of plastic to prevent distorting the genitalia. Photographs of genitalia were taken with a Leaf MicroLumina digital camera mounted on a Wild Photomakroskop M 400 stereo dissecting microscope. The images were enhanced with Adobe PhotoShop® 5.0 and imported into Adobe Illustrator® 6.0 to make the plates. Adult photo- graphs are from 35 mm slides that were made into digital images by a Polaroid SprintScan 35 slide scanner. These images were manipulated in Adobe PhotoShop® 5.0 and imported into Adobe Illustrator® 6.0 to make the plates. MORPHOLOGICAL TERMS General adult morphology, wing venation, and forewing pattern follows the nomenclature of Davis (1989), Miller (1991), and Lafontaine (1998). Genitalic nomenclature is compiled from several sources based on my ideas of homology of genital- ic parts within Spodoptera, especially the nomen- clature concerning the male valve. Terminology of the male tegumen and its attached parts (exclud- ing the valve) and the female genitalia follow Klots (1970) and Lafontaine (1998). The valve pre- sented problems of homology between species of Spodoptera. The structures of the valve follow the terminology of Forbes (1954) and are shown in Figs. 304-305. There is still confusion in the litera- ture concerning the structures of the valve. Table 1 lists different names used for homologous struc- tures of the valve from various authors. The costa is the sclerotized dorsoproximal margin of the valve. The cucullus is the dorsoapical region of the valve and is distinct ventrally from the valvula ex- cept in S. exigua. The valvula is the ventrodistal portion of the valve and the apex is distinct, except in S. exigua. The anellifer is the membranous re- gion in the central portion of the valve (Sibitani et al. 1954). In the central part of the inner surface of the valve there is a basal sclerite to which a muscle is attached to its dorsoproximal apex. This muscle controls the movement of the valve during copu- lation. In Spodoptera the basal sclerite is a plate with an oblique bar that can be variously modified (see character 12). From the basal sclerite arise two structures, the clasper proper and the ampulla. These 3 structures (basal sclerite, clasper proper, and ampulla) form the clasper. The clasper proper is the ventral projecting structure that lacks setae (Fig. 304). The clasper proper is present in clade 21 and is absent or reduced to a small dorsally pro- jecting flap in the rest of Spodoptera. The ampulla is always setose and is the prominent elongate pointed process in the central sub-apical portion of the valve (Figs. 304-305). An exception is in S. exempta, where the clasper proper is the large elon- gate pointed process, and the ampulla is a minute setose bump at the base of the clasper proper (Fig. 62). The editum is a prominent area of rather large setae just dorsal to the apex of the sacculus and sit- ting in the anellifer. It can be a small patch (Fig. 362) or be quite conspicuous (Fig. 360). Near the basal portion of the costa is a variously modified setose structure that ranges from a small tubercle (Figs. 41, 45) to an elongate process (Figs. 39, 43— 44), which I refer to simply as the costal process. This structure seems to be unique to Spodoptera. Bayer (1960) referred to this structure as the labis. The sacculus is the ventroproximal region of the valve and is usually more sclerotized than the valvula. Larval terminology follows that of Stehr (1987) and setal nomenclature follows Hinton (1946). Describing the color of various body parts and wing patterns in Lepidoptera is difficult because these colors are usually a mosaic of individual scales of many different colors. When describing color I will use the dominant color of the individ- ual scales making up that body part or wing pat- 8 REVISION OF SPODOPTERA GUENEE Table 1. Comparison of terms used in the male valve. The homologous structure are terms used in the Spodoptera valve and follow Forbes (1954). The costal process is first used here (see text). Homologous Reference Structure Structure Sibitani, et al. (1954) costa costa - editum sacculus sacculus valvula valvula cucullus cucullus harpe basal sclerite ampulla ampulla harpe clasper - costal process Klots (1970) costa costa editum editum sacculus sacculus valvula valvula cucullus cucullus - basal sclerite ampulla ampulla clasper clasper - costal process Todd and Poole (1980) costa costa = editum - sacculus valvula valvula cucullus cucullus - basal sclerite clasper ampulla 2 clasper ampulla costal process tern. If two or more colors are of approximately equal distribution the description will include each color. Individual moths within a species also show subtle variations in amount and intensity of colors. By describing only the dominant color, lengthy and confusing color descriptions can be avoided. Major variation in wing color and/or pattern will be discussed after each species de- scription. CLADISTIC ANALYSIS The taxa analyzed included 30 species of Spo- doptera and two outgroup taxa, Galgula partita Guenée and Elaphria grata Hiibner. The outgroups were selected based on a noctuid phylogeny de- Homologous Reference Structure Structure Lafontaine (1987) - costa - editum sacculus sacculus - valvula cucullus cucullus - basal sclerite harpe ampulla = clasper - costal process Lafontaine and - costa Poole (1991) - editum sacculus sacculus - valvula - cucullus - basal sclerite clasper ampulla = clasper = costal process Lafontaine (1998) - costa - editum sacculus sacculus - valvula - cucullus - basal sclerite clasper ampulla = clasper costal process rived from molecular data (Mitchell 1998). Male genitalia of G. partita are illustrated in Figs. 307 and 368 and the genitalia of E. grata are illustrated in Figs. 306 and 369. Female genitalia of G. partita are illustrated in Figs. 401 and 433 and the geni- talia of E. grata are illustrated in Figs. 400 and 432. These outgroups were used to test the monophyly of Spodoptera. A total of 24 characters was used of which 6 were binary and 18 multistate. Character state distributions of the multistate characters were as follows: three characters with 3 states, three with 4, two with 5 states, two with 6 states, three with 9 states, three with 10 states and two with 11 states. Fifteen multistate characters were coded as char- acter state trees (Characters 3-4, 7-13, 17-20, 22-23) (Figs. 4-17) (Mickevich 1982, Mickevich & M. G. POGUE 9 Weller 1990, Pogue & Mickevich 1990). Ordered multistate characters included 1 and 5-6. Data were analyzed using the Hennig86 parsimony program written by Farris (1988, Version 1.5). Missing data affects the placement of the taxa that contain missing data on the tree, resulting in multiple trees. Four vesica characters (19-22) were not available for two taxa, S. compta (Walker) and S. pectinicornis. The missing data were elimi- nated by inferring character states based on the states of their nearest neighbors. Transformation series analysis (TSA) was used to construct character state trees of the multistate characters (Mickevich 1982). TSA requires that character states of a multistate character are as- signed to each taxon on an initial cladogram. The initial cladogram of this analysis was created by treating all multistate characters as unordered. This will be referred to as the unordered tree(s). The trace character option in MacClade, Version 3 (Maddison & Maddison 1992) was useful in ana- lyzing character state change on a cladogram. By tracing character state change on a cladogram, character state trees can be developed (see Micke- vich & Weller 1990 and Pogue & Mickevich 1990 for methods of character state tree development). Most character state trees are complicated and have multiple branches (Figs. 6, 9-10, 15-17), and others are more simple (Figs. 4-5, 7, 11-13). Hen- nig86 requires linear variables, so the character state trees must be transformed into linear vari- ables. The method of transcribing non-linear character state trees into linear variables is termed ordinal coding (Mickevich & Weller 1990). For ex- ample, character 3 has five states (Fig. 4) with two state paths, the longest path consisting of four states (1-0-2-4), the shortest with two states (0-3). Ordinal coding produces two linear characters for each character state, one linear character defining the character state position on each of the paths. For character 3 (Fig. 4), state 1 is coded as 00 as the linear variable, state 0 is 10, state 2 is 20, state 4 is 30, and state 3 is 11. Since character state trees are ordered by how character states evolve on the cladogram, no assumption should be made about state “0” being plesiomorphic. More complicated character state trees with more paths will be rep- resented by more than two linear characters. The untransformed or raw data are in Appendix I and the transformed data are in Appendix II. The un- transformed data were used in the unordered analysis and the transformed data were used to produce the Spodoptera phylogeny. Character state trees are illustrated in Figs. 4-17. In some instances when analyzing the charac- ter state trees more than one equally parsimo- nious character state tree could be produced. I found that if you ran each of the character state trees individually with the rest of the data, one of the character state trees would usually produce a shorter tree. If there was more than one choice of character state tree, | would use the one that pro- duced the shortest tree. If all character state trees produced the same length tree, I chose the charac- ter state tree with the fewest paths. To find the shortest trees the “mhennig*” op- tion was used which constructs several trees, each by a single pass through the data and adding the taxa in a different sequence each time, and finally applies branch-swapping to each of the trees. The command “bb*” was then used which applies branch-swapping to each initial tree created by “mhennig*”, but retains all trees up to the limit of space allocated by the computer. CLADISTIC RESULTS The raw or untransformed data (Appendix I) were analyzed by treating all of the characters as unordered. This produced ten equally parsimo- nious trees with a length of 136 steps, a consisten- cy index (CI) of 0.79 and a retention index (RI) of 0.89. This step was necessary to produce a clado- gram on which to trace the character states of multistate characters in order to develop charac- ter state trees using TSA. The ten trees were iden- tical except in the clades on the cladograms desig- nated A-E (Figs. 1-2). Positional subtrees (Pogue & Mickevich 1990) were used to show the changes among the ten trees in Figs. 1-2. The unordered trees had two distinct tree topologies. Each one is represented in Figs. 1-2. The tree numbers are those produced by Hen- nig86, plus one, so Tree 0 in Henning86 will be Tree 1, Tree 1 is Tree 2, etc. in Table 2. Five areas (A-E) on the trees were changing, resulting in the 10 trees. In Fig. 1, four areas (B—E) were changing to produce the four trees represented in Table 2. Tree 1, with positional subtree cladograms, A1, B1, C1, D1, E1 is the tree shown in Fig. 1. To inter- pret Tree 6, positional subtree cladograms B2, C3, and E2 would be substituted for B1, C1, and E1; Tree 8 would have B2, C3, and D3 substituted on Fig. 1; Tree 10 would have B2 and C3 substituted 10 REVISION OF SPODOPTERA GUENEE descoinsi praefica cosmiodes ornithogalli latifascia marima evanida roseae E2 pulchella pulchella roseae E1 descoinsi marima cosmiodes ornithogalli evanida praefica latifascia littoralis dolichos litura androgea D3 | D1 m picta littoralis androgea litura dolichos picta frugiperda eridania compta C3 C1 ie compta eridania albula ee: albula ochrea ochrea triturata mauritia exempta pectinicornis malagasy B2 malagasy B1 apertura apertura pectinicornis umbraculata A1 pecten ie. cilium depravata exigua Elaphria Galgula Fig. 1. Phylogeny of Spodoptera produced by Hennig86 using 30 species and 2 outgroups (Elaphria and Galgula) with Elaphria designated as the outgroup. Twenty-four characters were used all coded as unordered (Appendix I). Ten equally parsimonious trees were constructed (length = 136 steps, CI = 0.79, RI = 0.89). Positional subtrees were used to define 4 of the 10 trees (see text and Table 2). M. G. POGUE praefica ornithogalli marima roseae pulchella E1 descoinsi cosmiodes evanida latifascia dolichos androgea D1 2 littoralis litura picta littoralis litura D3 picta androgea dolichos compta eridania ie albula ochrea C2 B2 A2 11 descoinsi cosmiodes latifascia evanida pulchella roseae marima ornithogalli praefica androgea dolichos littoralis litura picta frugiperda compta eridania albula ochrea triturata mauritia exempta pectinicornis malagasy apertura umbraculata cilium pecten depravata exigua Elaphria Galgula Fig. 2. Phylogeny of Spodoptera produced by Hennig86 using 30 species and 2 outgroups (Elaphria and Galgula) with Elaphria designated as the outgroup. Twenty-four characters were used all coded as unordered (Appendix I). Positional subtrees were used to define 6 of the 10 trees (see text and Table 2). 12 REVISION OF SPODOPTERA GUENEE on Fig. 1. The tree topology with positional sub- tree cladogram A2 are Trees 2-5, 7, and 9 from the Hennig86 output; Tree 5 is shown in Fig. 2. Hennig86 trees 2-4, 7, and 9 can be interpreted as above by substituting the positional subtree cladograms in Table 2 on Fig. 2. Using character state trees (Figs. 4-17), and transforming the character states to linear form us- ing ordinal coding (Mickevich & Weller 1990), pro- duced the data matrix in Appendix II. This data matrix was then analyzed, resulting in the final phylogeny shown in Fig. 3. Two trees were con- structed with a length of 146 steps, a CI of 0.76, and a RI of 0.90. The praefica species group had two dif- ferent topologies shown by the positional subtrees Al and A2, which resulted in the two trees. Monophyly of Spodoptera is supported by the synapomorphy of a dense cornutal patch on the vesica of the male aedoeagus and the female eighth segment covered with fine dense scales. The cladistic analysis of Spodoptera resulted in 8 monophyletic groups, with S. exigua the most plesiomorphic species. The cilium, umbraculata, and exempta species groups are Old World in ori- gin and whose larval host plants are primarily grasses. The larvae have a chisellike cutting sur- face to the mandible as compared to a serrate or toothed cutting surface found in the remaining Spodoptera larvae. The exception is S. pectinicornis whose larvae have serrate mandibles. The cilium Table 2. Ten unordered trees produced by Hen- nig86 using mhennig* option. Tree structure anal- ysis defined by positional subtree cladograms us- ing Figs. 2-3 (see text). Tree Fig. number Positional subtree cladograms number 1 Als Biel (Di 2 EI 1 6 ANY B25 sE3; D1 E2 1 8 Alls 1B2:-.G3) D3 "El 1 10 Aly BZ E3- Dil El 1 2: AQs (B2s C2 Die El 2 3 AD BZ puG20 192) oi 2 4 AO Bl Cleese) El 2 5 AZ: GB2 5 8C25 D2 4E2 2 Uf A2) B27 2Cl) (Die? 2 9 A220 SZ Glo Dig vb 2 species group is defined by 4 characters (Appen- dix III, Clade 25). The umbraculata species group is defined by 2 male genitalic characters (Appen- dix III, Clade 27). The exempta species group is defined by a male genitalic character (Appendix III, Clade 18). Clade 21 is of Neotropical origin with S. eridania and S. albula extending into the Nearctic. This species group is defined by 6 char- acters (Appendix III, Clade 21) 5 from the male genitalia and 1 from the female genitalia. Spodop- tera frugiperda is the sister species to the remain- ing species groups (dolichos, littoralis, evanida, and praefica). These groups have a similar forewing pattern (Figs. 253-303) with the M vein of the forewing being highlighted. All are New World in origin except the littoralis clade. The dolichos species group is defined by 3 male genitalic char- acters (Appendix III, Clade 17). The littoralis species group is defined by 5 male genitalic char- acters (Appendix III, Clade 15). The evanida species group is defined by 6 male genitalic char- acters (Appendix III, Clade 12). The praefica species group is defined by four male genitalic characters and one female genitalic character (Appendix III, Clade 9). There are several species pairs of Spodoptera that were either considered the same or very closely related species. All of these pairs are sister species on the cladogram. These include S. apertu- ra and S. malagasy which were originally de- scribed as subspecies, S. littoralis and S. litura were considered conspecific, as were S. latifascia and S. cosmiodes. The genitalia are virtually the same in S. ornithogalli and S. marima, but S. marima is not sexually dimorphic and is geographically isolated from S. ornithogalli. This is reflected in Fig. 3, subtree Al as S. marima and S. ornithogalli form a trifurcation with Clade 11. PHYLOGENETIC CHARACTERS The following is a list of the characters used in reconstructing the phylogeny of Spodoptera. They are arranged in order of tagma. Binary characters have the plesiomorphic state coded as 0 and de- rived state as 1. Multistate characters are coded us- ing ordinal coding (Mickevich & Weller 1990). Character state trees are illustrated for fourteen multistate characters (Figs 4-17). Two sets of char- acter states are referred to. The states in bold are the raw data, presented in Appendix I.The states in italics are the transformed data coded ordinally M. G. POGUE 12 ornithogalli 11 marima pulchella A2 a roseae praefica 20 12 21 25 ilk 10 28 14 16 17 19 13 roseae pulchella ornithogalli marima praefica evanida latifascia cosmiodes descoinsi littoralis litura picta androgea dolichos frugiperda exempta mauritia triturata albula ochrea eridania compta pecten depravata cilium umbraculata pectinicornis apertura malagasy exigua Elaphria Galgula Fig. 3. Phylogeny of Spodoptera produced by Hennig86 using 30 species and 2 outgroups (Galgula and Elaphria) with Galgula designated as the outgroup. Twenty-four characters were used consisting of 6 binary and 18 multi- state. Multistate characters were coded as 3 ordered and 15 defined by cladogram characters presented in Figs. 4-17 and in Appendix II. Two equally parsimonius trees were constructed (length = 143 steps, CI = 0.77, RI = 0.91) Positional subtrees Al and A2 were used to define these trees. REVISION OF SPODOPTERA GUENEE (3) (4) D-O-2-& 030,050 (8) oEOOo . @) © i 2 Q @® 0-050-0 O-M-@BHO-D © B) 8 ) 9 Q Oe Y G-0-0 OM2H-O ® @ 10 11 Figs. 4-11. Character state trees produced using TSA (see text) and used in data matrix in Appendix II. 4, Character 3.5, Character 4. 6, Character 7. 7, Character 8. 8, Character 9. 9, Character 10. 10, Character 11. 11, Character 12. to reflect the relationships in the character state have a filiform antenna which is considered ple- trees and are presented in Appendix II. siomorphic. Bipectinate antennae arose twice, in Clade 24 and in S. roseae (Fig. 3). It is possible that H the bipectination in Clade 24 is not homologous ead ‘ ; with that in S. roseae. Serrate antennae arose twice, Antenna: Character 1. Form of male flagellum. (0) in S. cilium and in Clade 29 (S. apertura and S. (0): filiform (Fig. 18); (1) (1): bipectinate (Fig. 19); malagasy). The character was treated as ordered. (2) (2): serrate (Fig. 20). Most species of Spodoptera CI = 0.50 oe Q\ /® oN ©) M. G. POGUE O50,050,0,050 ole ODO @-8)- 16 14 15 Ota S-@ 8) @ 17 Figs. 12-17. Character state trees produced using TSA (see text) and used in data matrix in Appendix II. 12, Char- acter 13. 13, Characters 17 and 22. 14, Character 18. 15, Character 19. 16, Character 20. 17, Character 23. Thorax Prothoracic tibia: Character 2. Length of lateral scale tufts. (0) (0): not extending beyond first tarsal segment (Fig. 21); (1) (1): extending beyond first tarsal segment (Fig. 22). The scales of the protho- racic tibia are elongate and extend beyond the first tarsal segment in S. mauritia and S. triturata, which is a good character to separate them from the rest of Spodoptera. CI = 1.00. Mesothoracic tibia: Character 3. Tibial spur scal- ing. (0) (20): spurs completely scaled (Fig. 23); (1) (10): mediolateral surface broadly devoid of scales on both spurs (Fig. 24); (2) (30): mediolater- al surface thinly devoid of scales on both spurs (Fig. 25); (3) (21): mediolateral surface devoid of 16 REVISION OF SPODOPTERA GUENEE Figs. 18-20. Antenna: Character 1, form of male flagellum. 18, Filiform, state 0, S. exigua. 19, Bipectinate, state 1, S. pectum. 20, Serrate, state 2, S. cilium. Figs. 21-22. Prothoracic tibia: Character 2, length of lateral scale tufts. 21, Not extending beyond first tarsal segment, state 0, S. eridania. 22, Extending beyond first tarsal segment, state 1, S. mau- ritia. scales on large spur (Fig. 26); (4) (00): internal sur- face at base thinly devoid of scales on large spur (Fig. 27). The lack of scales on the male mesotho- racic tibial spurs of some species was first men- tioned by Zimmerman (1958). Spurs can be com- pletely scaled or have varying degrees of bare ar- eas on one or both spurs. Viette (1967) and Brown & Dewhurst (1975) used this character in keys to Madagascan and African Spodoptera. Character state tree - Fig. 4. CI = 0.50. M. G. POGUE 17 Figs. 23-27. Mesothoracic tibia: Character 3, tibial spur scaling. 23, Spurs completely scaled, state 0, S. eridania. 24, Medio-lateral surface broadly devoid of scales on both spurs, state 1, S. mauritia. 25, Medio-lateral surface thinly devoid of scales on both spurs, state 2, S. exigua. 26, Medio-lateral surface devoid of scales on large spur, state 3, S. albula. 27, Internal surface at base thinly devoid of scales on large spur, state 4, S. roseae. Male Genitalia apex (Fig. 30); (3) (30): narrowest at base, widest distally (Fig. 31). (4) (21): very wide, gradually Uncus: Character 4. Width. (0) (10): wide, con- narrowing to apex (Fig. 32). Most Spodoptera have stant throughout length (Fig. 28); (1) (00): narrow an uncus that is widest at base and gradually nar- at base, wide medially to produced apex (Fig. 29); rowing to apex. Some Old World species have de- (2) (20): widest at base, gradually narrowing to veloped an uncus with a wide distal half, which 18 REVISION OF SPODOPTERA GUENEE arose on Clades 19 and 25 (Fig. 3). Spodoptera um- braculata has a unique uncus shape that is very wide and gradually narrows to its apex (Fig. 32). Character state tree - Fig. 5. CI = 0.80. Scaphium: Character 5. Form and amount of scle- rotization. (0) (0): absent (Fig. 306); (1) (1): elon- gate, weakly developed (Fig. 33); (2) (2): elongate, well sclerotized (Fig. 34). The tuba analis is the posterior part of the digestive tract and is sup- ported by the dorsal scaphium (Klots 1970). In Clade 21 the scaphium is well sclerotized and elongate. This character was treated as ordered, with (0) as the plesiomorphic state. CI = 1.00. Tegumen: Character 6. Lateral projections of tegumen. (0) (0): lacking a pair of projecting arms on upper third (Fig. 35); (1) (1): lateral swellings at basal two-thirds (Fig. 36); (2) (2): with a pair of projecting arms on upper third (Fig. 37). In Clade 21 the lateral edge of the tegumen is modified from the state found in other Spodoptera. It is swollen in S. compta and S. ochrea and has promi- nent projecting arms in S. eridania and S. albula. This character was treated as ordered, with 0 as the plesiomorphic state. CI = 0.67. Costal process: Character 7. Shape. (0)(011): ab- sent (Fig. 38); (1) (111): elongate, narrow (Fig. 39); (2) (311): short, straight, narrow (Fig. 40); (3) (211): a minute knob (Fig. 41); (4) (110): short, wide (Fig. 42); (5) (112): elongate, wide, slightly curved (Fig. 43); (6) (121): elongate, wide, well sclerotized (Fig. 44); (7) (411): a small swelling, no distinct process (Fig. 45); (8) (101): medium length and width (Fig. 46). The costal process is located sub- basally on the costa of the valve. It is found in all species of Spodoptera, except S. exigua, where it may have been secondarily lost. It has many forms, from a minute knob (Fig. 41) or small swelling (Fig. 45) to an elongate, well sclerotized process (Fig. 44). In Todd & Poole (1980) this structure was misinterpreted as the ampulla. Character state tree - Fig. 6. CI = 0.80. Character 8. Location on costa. (0) (00): absent (Fig. 47); (1) (10): at base of costa on inner edge (Fig. 48); (2) (20): in middle of costa (Fig. 49); (3) (11): at base of costa along margin (Fig. 50). The costal process can be located on the margin, middle, or inner edge of the costa. Character state tree - Fig. 7. CI = 1.00. Cucullus: Character 9. Form. (0) (11): distinct from valve (Fig. 51); (1) (01): pointed (Fig. 52); (2) (20):broadly rounded (Fig. 53); (3) (31): curved at costal apex, outer margin straight (Fig. 54); (4) (21): evenly rounded from costal apex to valvula apex (Fig. 55); (5) (22): truncate, forming almost a right angle at costal apex, outer margin straight (Fig. 56). The cucullus is the dorsodistal area of the valve. The outer margin varies from broadly rounded to truncate. Character state tree - Fig. 8. CI = 0.63. Ampulla: Character 10. Form. (0) (0000): elon- gate, sinuate process, apex pointed (Fig. 57); (1) (1000): elongate process bent at a right angle at 2/3 length (Fig. 58); (2) (2000): elongate, narrow, slightly curved, apex round (Fig. 59); (3) (3100): short, thumb-shaped lobe, apex round (Fig. 60); (4) (3200): short, triangular shaped lobe, apex round (Fig. 61); (5) (3010): minute lobe (Fig. 62); (6) (4000): elongate, slightly curved with a point- ed produced apex (Fig. 63); (7) (5000): elongate, slightly curved with an angulate pointed apex (Fig. 64); (8) (3000): elongate, slightly curved with a decurved apex (Fig. 65); (9) (3001): elongate, broad, curved with a decurved apex (Fig. 66). The ampulla is a structure that has been confused in the literature (Table 1). The ampulla is part of the clasper in the broad sense (Forbes 1954) and is dorsodistal from the basal sclerite (Figs. 67-76) to which a muscle is attached (Fig. 70, m). The am- pulla is the transverse costal process with fine sensory setae and the sclerotized ventrodistal process is the clasper proper (Figs. 60-63, amp, cl). In most species of Spodoptera only the ampulla is present. In S. exempta the ampulla is a small knob located at the base of a well developed clasper (Fig. 62). Character state tree - Fig. 9. CI = 1.00. Basal sclerite of clasper: Character 11. Form. (0)(0110): short, crescent-shaped (Fig. 67); (1) (1010): oblique bar (Fig. 68); (2) (1110): a narrow plate with an oblique bar at proximal end (Fig. 69); (3) (1111): an elongate plate with costal and oblique bars joined to form a horizontal X (Fig. 70); (4) (1210): a broad plate with an oblique bar at proximal end (Fig. 71); (5) (2100): a quadrate plate with slightly sclerotized costal bar and an elon- gate horizontal bar (Fig. 72); (6) (2110): plate re- duced, costal and oblique bars form an obtuse an- gle (Fig. 73); (7) (2120): an oblong plate, costal and oblique bars form an acute angle (Fig. 74); (8) (3110): a rectangular plate with costal and oblique bars joined to form a horizontal X (Fig. 75); (9) (4110): a reduced plate, costal bar curved and join- ing oblique bar to form an inverted Y (Fig. 76). The basal sclerite of the clasper, in the broad M. G. POGUE 19 33 34 Figs. 28-32. Uncus: Character 4, width. 28, Wide, constant throughout length, state 0, Galgula partita Guenée. 29, Narrow at base, wide medially to produced apex, state 1, Elaphria grata Hiibner. 30, Widest at base, gradually nar- rowing to apex, state 2, S. albula. 31, Narrowest at base, widest distally, state 3, S. mauritia. 32, Very wide, gradual- ly narrowing to pointed apex, state 4, S. umbraculata. Figs. 33-34. Scaphium: Character 5, form and amount of scle- rotization. 33, Elongate, weakly developed, state 1, S. evanida. 34, Elongate, well sclerotized, state 2, S. eridania. 20 REVISION OF SPODOPTERA GUENEE Sif Figs. 35-37. Tegumen: Character 6, lateral projections of tegumen. 35, Lacking a pair of projecting arms on upper third, state 0, S. ornithogalli. 36, Lateral swellings at basal two-thirds, state 1, S. ochrea. 37, Pair of projecting arms on upper third, state 2, S. eridania. sense, is defined as the sclerite to which the valve muscle is attached. In Spodoptera the point of at- tachment is a small dorsocostal process (Fig. 70, m). Character state tree - Fig. 10. CI = 0.90. Clavus: Character 12. Shape. (0) (01): a large, spiny, ellipitical lobe (Fig. 307); (1) (11): absent (Fig. 77); (2) (21): a minute round projection (Fig. 78); (3) (20): elongate, finger-shaped (Fig. 79); (4) (31): large, club-shaped, base narrower than apex (Fig. 80); (5) (41): short, round, covered with setae (Fig. 81). The clavus is a small separate organ be- tween the base of the valve and juxta bearing sen- sory setae. It is absent in the more plesiomorphic Spodoptera, except in S. pectinicornis, S. apertura, and S. malagasy, and S. mauritia where it is a minute round projection. This character state is also shared with S. frugiperda and the littoralis species group. Character state tree - Fig. 11. Cl = 0.71. Sacculus: Character 13. Shape. (0)(01111): cylin- drical with heavily scelerotized triangular apex (Fig. 82); (1) (10111): quadrate (Fig. 83); (2) (12111): M. G. POGUE 21 Figs. 38-46. Costal process: Character 7, shape. 38, Absent, state 0, S. exigua. 39, Elongate, narrow, state 1, S. albula. 40, Short, straight, narrow, state 2, S. triturata. 41, A minute knob, state 3, S. cilium. 42, Short, wide, state 4, S. erida- nia. 43, Elongate, wide, slightly curved,state 5, S. picta. 44, Elongate, wide, well sclerotized, state 6, S. latifascia. 45, A small swelling, no distinct process,state 7, S. exempta. 46, Medium length and width, state 8, S. praefica. 22 REVISION OF SPODOPTERA GUENEE 90 Figs. 47-50. Costal process: Character 8, location on costa. 47, Absent, state 0, S. exigua. 48, At base of costa on in- ner edge, state 1, S. picta. 49, In middle of costa, state 2, S. triturata. 50, At base of costa along margin, state 3, S. mauritia. broad to half its length, tapering distally (Fig. 84). (3) (11111): rectangular, proximal margin wider than distal (Fig. 85); (4) (21111): trapezoidal, dor- sal margin angulate (Fig. 86); (5) (11011): elongate, basal notch, widest at one-quarter length, gradu- ally tapered to apex (Fig. 87); (6) (11211): elongate, basal notch, widest at one-third length, abruptly tapered to apex (Fig. 88); (7) (11101): triangulate (Fig. 89); (8) (11121): rectangular, slightly con- stricted at two-thirds length, apex indented (Fig. 90); (9) (11110): quadrate, apex tapered distally (Fig. 91); (10) (11112): rectangular, apex truncate dorsally, an elongate taper ventrally (Fig. 92). The sacculus is the ventrobasal portion of the valve which is more sclerotized ventrally and more membranous dorsodistally where it is clothed with setae. Various forms are present that define the different clades (Appendix III). Character state tree - Fig. 12. CI = 1.00. Valvula: Character 14. Ventral margin shape. (0)(0): smooth, setae not originating on short pro- tuberances (Fig. 93); (1) (1): scalloped, setae origi- nating on short protuberances (Fig. 94). The ven- tral margin of the valvula can be smooth or scalloped with setae arising on short protuber- ances. The scalloped edge is found in S. frugiperda and all of its sister taxa (Fig. 3). CI = 1.00. Character 15. Width. (0)(0): narrower than cu- cullus (Fig. 93); (1) (1): wider than cucullus (Fig. 94). The width of the valve also divides Spodoptera into S. frugiperda and all of its sister taxa. CI = 0.33. Character 16. Valvular indentation. (0) (0): ab- sent (Fig. 95); (1) (1): present (Fig. 96). Bayer (1960) recognized the valvular indentation just M. G. POGUE 23 Figs. 51-56. Cucullus: Character 9, form. 51, Distinct from valve, state 0, G. partita. 52, Pointed, state 1, E. grata. 53, Broadly rounded, state 2, S. albula. 54, Curved at costal apex, outer margin straight, state 3, S. androgea. 55, Evenly rounded from costal apex to valvula apex, state 4, S. mauritia. 56, Truncate, forming almost a right angle at costal apex, outer margin straight, state 5, S. frugiperda. below the juncture of the basal sclerite and saccu- lus. It is present in S. frugiperda and all of its sister taxa. CI = 1.00. Coremata: Character 17. Number of lobes. (0)(00): absent (Fig. 97); (1) (20): no distinct lobe, ap- pressed against sacculus (Fig. 98); (2) (10): single lobe (Fig. 99); (3) (11): double lobe (Fig. 100). Core- mata are large lobes covered with long fine hairs that are attached basally to the dorsal region of the valve. The coremata disperse pheromones during mating. All Spodoptera have coremata with varying degrees of development. In the cilium and umbraculata clades the coremata are reduced and closely appressed to the valvula. In the remaining species of Spodoptera the coremata can be either a single or in form of a double lobe. Character state tree— Biss 13: 1Gl = (0160: Juxta: Character 18. Shape. (0)(11): hexagonal plate with dorsolateral arms (Fig. 101); (1) (01): helmet-shaped (Fig. 102); (2) (33): a narrow rec- tangular band with ventrolateral projections (Fig. 103); (3) (32): a rectangular band, median process elongate, expanding into an apical plate (Fig. 104); (4) (21): elliptical (Fig. 105); (5) (31): a narrow band, median process triangulate (Fig. 106); (6) (30): quadrate with lateral third distinct (Fig. 107); (7) (41): a narrow band expanded ventrolaterally, median projection broad, triangular (Fig. 108); (8) (51): a narrow band expanded ventrolaterally, me- dian projection elongate, triangular (Fig. 109); (9) (61): anarrow band expanded ventrolaterally, me- dian projection an elongate narrow triangle (Fig. 110). The juxta is a flat plate that joins the bases of the valves. Character state tree - Fig. 14. CI = 0.90. Vesica: Character 19. Dorsobasal sclerotized patch. (0)(110): absent (Fig. 111); (1) (220): quadrate, gran- 24 REVISION OF SPODOPTERA GUENEE 66 Figs. 57-66. Ampullla: Character 10, form. 57, Elongate, sinuate process, apex pointed, state 0, G. partita. 58, Elon- gate process bent at a right angle at 2/3 length, state 1, E. grata. 59, Elongate, narrow, slightly curved, apex round, state 2, S. exigua. 60, Short, thumb-shaped lobe, apex round, state 3, S. albula. 61, Short, triangular shaped lobe, apex round, state 4, S. ochrea. 62, Minute lobe, state 5, S. exempta. 63, Elongate, slightly curved with a pointed pro- duced apex, state 6, S. cilium. 64, Elongate, slightly curved with an angulate pointed apex, state 7, S. apertura. 65, Elongate, slightly curved with a decurved apex, state 8, S. androgea. 66, Elongate, broad, curved with a decurved apex, state 9, S. latifascia. M. G. POGUE 25 is Figs. 67-76. Basal sclerite of clasper: Character 11, form. 67, Short, crescent-shaped, state 0, G. partita. 68, Oblique bar, state 1, S. exigua. 69, A narrow plate with an oblique bar at proximal end, state 2, S. exempta. 70, An elongate plate with costal and oblique bars joined to form a horizontal X, state 3, S. apertura. 71, A broad plate with an oblique bar at proximal end, state 4, S. albula. 72, A quadrate plate with slightly sclerotized costal bar and an elon- gate horizontal bar, state 5, S. androgea. 73, Plate reduced, costal and oblique bars form an obtuse angle, state 6, S. frugiperda. 74, An oblong plate, costal and oblique bars form an acute angle, state 7, S. litura. 75, A rectangular plate with costal and oblique bars joined to form a horizontal X, state 8, S. pulchella. 76, A reduced plate, costal bar curved and joining oblique bar to form an inverted Y, state 9, S. cosmiodes. 26 REVISION OF SPODOPTERA GUENEE 81 Figs. 77-81. Clavus: Character 12, shape. 77, Absent, state 1, S. exigua. 78, A minute round projection, state 2, S. apertura. 79, Elongate, finger-shaped, state 3, S. dolichos. 80, Large, club-shaped, base narrower than apex, state 4, S. latifascia. 81, Short, round, covered with setae, state 5, S. ornithogalli. ulose (Fig. 112); (2) (200): single elongate spine (Fig. 113); (3) (210): narrow, elongate, aculeate (Fig. 114); (4) (211): dense patch of elongate spines (Fig. 115); (5) (010): narrow, elongate, granulose (Fig. 116); (6) (310): wide, elongate, aculeate (Fig. 117); (7) (510): narrow, elongate, granulose basally, aculeate apically (Fig. 118); (8) (33): quadrate, ac- uleate (Fig. 119). At the base of the vesica, where it emerges from the aedoeagus, is a variously shaped spiculate patch. The patch varies consider- ably from an indistinct patch with minute spicula- tions (Fig. 112) to a well developed spine (Fig. 113) or patch of large spines (Fig. 115). Character state tree - Fig. 15. CI = 0.89. Character 20. Distal cornutus. (0)(01000): conical (Fig. 120); (1) (11000): multiple conical cornuti (Fig. 121); (2) (41000): ovate, aculeate apically (Fig. 122); (3) (31000): absent (Fig. 123); (4) (21000): large spine (Fig. 124); (5) (20000): patch of large spines within dense cornutal patch (Fig. 125); (6) M. G. POGUE 27 Lf 4 7 Figs. 82-92. Sacculus: Character 13, shape. 82, Cylindrical with heavily scelerotized triangular apex, state 0, G. par- tita. 83, Quadrate, state 1, E. grata. 84, Broad to half its length, tapering distally, state 2, S. albula. 85, Rectangular, proximal margin wider than distal, state 3, S. exigua. 86, Trapezoidal, dorsal margin angulate, state 4, S. cilium. 87, Elongate, basal notch, widest at one-quarter length, gradually tapered to apex, state 5, S. litura . 88, Elongate, basal notch, widest at one-third length, abruptly tapered to apex, state 6, S. pulchella. 89,Triagulate, state 7, S. apertura. 90, Rectangular, slightly constricted at two-thirds length, apex indented, state 8, S. cosmiodes. 91, Quadrate, apex ta- pered distally, state 9, S. androgea. 92, Rectangular, apex truncate dorsally, an elongate taper ventrally, state 10, S. dolichos. 28 REVISION OF SPODOPTERA GUENEE og Figs. 93-96. Valvula characters. 93, Character 14: ventral margin shape: smooth, setae not originating on short pro- tuberances, state 0; character 15: narrower than cucullus (A>B), state 0, S. albula. 94, Character 14: ventral magrin shape: scalloped, setae originating on short protuberances, state 1; character 15: wider than cucullus (A se bee e oe eee eee aere S. triturata (Walker) 3 Forewing ground color cream with an irregular patch of rose scales from oblique ground color band to postmedian line between costa and R vein (Fig. 278); forewing length 18-22 mm; Oriental, Aus- tralasian,and' ©ceanial (Fig..038) |p siya sty oops re ae te eee: S. picta (Guérin-Meéneville) 5 2 Forewing ground color not.cream: and noirose'scales present )3.4.5-1-.0040¢ 9 eee eee 10 Forewing length greateritham UO mm) 23.7. 22g cine kits aie ie ee 11 Forewinevengthiless thane 0mm mat soils eer ioc S. pectinicornis (Hampson) @ Forewing with orbicular spot round (Figs. 172-180, 182, 184-185, 187-188, 206) .................... 12 Forewing with orbicular spot oval or absent (Figs. 193, 196, 198, 200, 202-203, 230-234, 236-240, 246-247, DAs) Va 252 Pe anaes ire arere e ete el aya eR RN rar At clint nema Ur a Rhemrar inert ote e e070 0-050.0.000000¢ 14 Forewing with orbicular spot an oblique trapezoid that extends from costal margin to Cu vein (Figs. 212, 2187263267; 2608=277)) sucans Ab eas Feta int eine eas ede eel crete I Se en 18 Hindwing with apex, outer margin, and veins infuscated with light brown scales (Figs. 172-180, 206) 13 Hindwing with gray scales along apex and outer margin, veins not infuscated with gray scales (Figs. 182, afey Sake loyal keV Goto) ame emcee Ne AHO arg Rene mig ocroldlo H Line oom ou mao oC c S. cilium Guenée ¢ Forewing orbicular spot large, cream, no contrasting color in center, faintly outlined in black; length greater than 1 Sammi (Rig7206)- sams neice eee OOo eee ene S. umbraculata (Walker) @ Forewing orbicular spot small, cream, center buff, not outlined in black; length less than 15 mm (Figs. 7D NBO in Bere Be cag spae pees ce aokcts sina ents MLO og Ne aa S. exigua (Hiibner) 3 & Mesothoracic tarsus fuscous with cream colored apical/rings) 00)... 5-5. oss el eee 15 Mesothoracic tarsus cream/colored) 2. Sos bel eis- Gie es seioe oe oe Oe ee eee 17 Hindwings with veins infuscated with brown to base (Figs. 230-234, 236-240, 246-247) ............. 16 Hindwings with veins infuscated with brown at apex, white to base (Figs. 250, 252) ................-- Re Ta ets cre ae cr ionT Aine ee Nitec IOI oid Hence o CCC oO. IB eo ecw Oo aS S. triturata (Walker) 2 Forewing orbicular spot distinct, with buff to brown center, in some specimens outlined in black; reniform spot dark surrounded by a white to cream ring and an outer black ring (Figs. 230-234) ee ae ONT Se Rat ts TAN a ON EN ARUN 1) eal Ce ROR OT BY Ne S. exempta (Walker) 3 2 Forewing orbicular spot faint to absent, never outlined in black; reniform spot black, lacking any border (Figs..236-240) 246-247) cs ctles sop cui temitns satel mule Seen leyortse ht nate ae eee S. mauritia 2 Forewing with very faint wing markings, antemedial and postmedial lines absent, orbicular spot very faint to absent, reniform spot round, black and faint; hindwing white with apex, outer margin to vein CuA2, and distal ends of veins Rs to CuA2 with light brown scales (Figs. 196, 198, 200, 202-203) Ee isleyalla ahacsleyrohtin ve ehcce Peer esp Se eele 5y ay aco era yew fe al ce Meteo te ON oe Steet eee ae S. pecten Guenée @ Forewing with wing markings present; hindwing white suffused with pale gray (Fig. 193) ............ PONT DO SG me a cra ears Men ue MINN aS roa as Cindi ol 6 Oe Ue bit.od o.0 abn's S. depravata (Butler) 2 Forewing with reniform spot brown outlined in white on proximal border, distal apex with a triangular light brown spot outlined in white, a white distal border extending to middle (Figs. 263-277) ..... 19 Forewing with reniform spot crescent-shaped, lacking light brown triangular spot outlined in white at apex (Figs..212, 218) in reatsgeiard sues HeieiOlue 6, omustsleyd sl cao theses sieht Een ees eee ene 20 Hindwing veins infuscated with brown scales apically (Figs. 268-277); Oriental, Australasia, and eastern Palaearctic regions: (Figs'537)- ese sans seine eee ie ee eens eT eer S. litura (Fabricius) 3 2 Hindwing veins not infuscated with brown scales near apex (Figs. 263-267); western Palaearctic and Ethiopian'regions| (Fig. 537). ase secereictace eet es elena S. littoralis (Boisduval) 3 @ Forewing ground color dark brown, reniform spot cream outlined in black, center light brown to brown; hindwangwihite (Fig; 212) 0. 5 o ve ec nie n soerrtio ae oe Mme nee ists cette S. apertura (Walker) 2 Forewing ground color brown, reniform spot cream outlined in black, center light brown with a white median line; hindwing light brown (Fig. 218); endemic to Madagascar (Fig. 528) .S. malagasy Viette ° KEY TO ADULTS OF NEW WORLD SPECIES Antenna serrate (Fig. 20); endemic to the Galapagos Islands (Fig. 543) ............. S. roseae (Schaus) 3 Antenna:filiformy (Figs:18) iii. 8 census Gia dat ae Ue eenO onan taPe St braid Orel ine uate eee 2 Forewing orbicular spot round) (Figs)172-180) 303) fiayscmieloe old. cere cialis aie ye ok: eee 3 ill(S)). i", 12(11’). 22 13(4’). IS 14(13). 14’. 15(14). iey 16(6’). 16’. 17(16). M. G. POGUE 39 Forewing with orbicular spot oval (Figs. 222, 225, 227-228) or an oblique trapezoid extending from costal mareinitoeulveim|(Eigsy2o9/261) 284-286-2877 290-2911) 293/296) S00). iarieery so rietdsalaeler yr -ie)eraiie 4 Forewing with orbicular spot cream, with pale rufous center; forewing length 11-14 mm (Figs. 172-180) eG Bere cba Go to'o.0'01a 6 Of .0 6/000 COPS CIOL CHOU CR ICT EERIE CERO CP acter ect cl ean S. exigua (Hiibner) 3 & Forewing with orbicular spot cream, with buff to brown center, outlined in black; forewing length 20-22 mm (Fig. 303); endemic to the Galapagos Islands (Fig. 543) .................55. S. roseae (Schaus) 2 Forewing with no white scales along Cu vein under orbicular and reniform spots (Figs. 219-229, 253-258, D999) SRST Se Ie IG Te EE hy Ses Pee ea eee w TIA D As Sle ROR Sel MEE coe alert 5 Forewing with white scales along Cu vein under orbicular and reniform spots extending to M3 and Cul veins, continuing down CuA2 vein to postmedial line (Figs. 259-262, 291-300) ................. 13 Eorewanesoroundi colon rufous (Figs 2192235283), 285-290) i, «cto ieia's daeieterciens tata oieyerieie severe oe 6 Horewaneserouncicol orpalerenayion cream) (Fies3224—229727 9-281) 5 5 seme ci stsisle se eieoelss stele ee 9 Forewing ground color brown, or dark brown (Figs. 253-258, 282, 284) ............... cece eee 11 Forewing with costal and posterior margin areas concolorous (Figs. 219-220, 279-281, 285-286) ...... Uf Forewing with costal area darker than posterior marginal area (Figs. 283, 287-290) ...............5. 16 Forewing with a distinct pale gray postmedian band between subterminal and adterminal lines (Figs. DOD = SO) watves aids aie ae ate rencnche Mactan ota a lceeee ns, aaxtial ape nialesats Sales S. cosmiodes (Walker) 3 Forewing lacking a distinct postmedian band between subterminal and adterminal lines (Figs. 219-220, TID SMW ie MA ENT ONE SPR ICES Sy Ley ENA Se Pots ee as FAV sis beh, USSa eae ote “hs “a geet ay : c = ‘ ‘ 7 at 125 Y if a rs i. ng | a fof in ( ia, | = Y } Onl he. ster : ; vA ) v) ‘_ 1 j ’ ( s j yh a ] / : N Ya e's uy { / f ) i rE he F im om Re fe = 1 { Rs ; F { 4 \ | \ ns / ne ) 7) | x ; : ' 4